by Allison Baker
figures by Lillian Horin

The Arctic apple is the juiciest newcomer to produce aisles. It has the special ability to resist browning after being cut (Figure 1), which protects its flavor and nutritional value. Browning also contributes to food waste by causing unappealing bruising on perfectly edible apples. Food waste, especially for fruits and vegetables, is a major problem worldwide; nearly half of the produce that’s grown in the United States is thrown away, and the UK supermarket Tesco estimates that consumer behavior significantly contributes to the 40% of its apples that are wasted. Therefore, Arctic apples not only make convenient snacks, but they also might be able to mitigate a major source of food waste.

Figure 1: Traditional Golden Delicious apple (left) versus the Arctic variety (right). After slicing into the apples, the traditional Golden Delicious apple is turning brown as expected. On the other hand, the Arctic Golden doesn’t become discolored at all. (Image credit: Okanagan Specialty Fruits Inc.)

While a non-browning apple sounds great, how exactly was this achieved? Arctic apples are genetically engineered (GE) to prevent browning. This means that the genetic material that dictates how the apple tree grows and develops was altered using biotechnology tools. But before learning about the modern science used to make Arctic apples, let’s explore how traditional apple varieties are grown.

How are new apple varieties developed?

Harvesting tasty apples is more complicated than simply planting a seed in the ground and waiting for a tree to grow. In particular, it’s difficult to predict what an apple grown from a seed will look and taste like because each seed contains a combination of genetic material from its parents. But farmers can reliably grow orchards of tasty apples by using an ancient technique called grafting. After a tree that produces a desirable apple is chosen, cuttings of that original tree are grafted, or fused, onto the already-established roots of a donor tree, called rootstock. The cuttings then grow into a full-sized tree that contains the exact same genetic material as the original tree. As a result, each tree of a specific apple variety is a cloned descendant of the original tree, and thus produce very similar apples.

New apple varieties emerge when genetic changes are allowed to occur. Traditionally, new apples are produced by cross-breeding existing apple varieties. This reshuffles the genetic makeup of seeds, which are then planted to see if they grow into trees that produce delicious new apples. On the other hand, Arctic apples are created by making a targeted change to the genetic material of an existing variety (more on this later). The advantage of using genetic engineering over traditional breeding methods is that scientists can efficiently make precise improvements to already-beloved apple varieties—in contrast, traditional cross-breeding is much more random and difficult to control.

What causes apple browning?

Insight into the molecular causes of apple browning guided the genetic alteration that made Arctic apples. Apples naturally contain chemicals known as polyphenols that can react with oxygen in the air to cause browning. This reaction won’t occur, however, without the help of polyphenol oxidase (PPO) enzymes, which bring polyphenols and oxygen together in just the right way. PPO enzymes and polyphenols are normally separated into different compartments in apple cells, which is why the inside of a fresh apple is white or slightly yellow-green in color. But these structures are broken when the fruit is cut or crushed, allowing PPOs to interact with polyphenols and oxygen to drive the browning reaction (Figure 2). This process occurs in all apples, but some varieties are less susceptible than others due to factors like lower amounts of PPOs or polyphenols. Common household tricks can also delay browning by a few hours by interfering with the PPO reaction, but no method prevents it completely or indefinitely. Knowing that PPOs were responsible for browning, researchers thought about blocking the production of these enzymes with genetic tools to create non-browning apples.

Figure 2: The molecular-level science behind Arctic Apples. PPO enzymes (red ovals), polyphenols (purple hexagons), and oxygen (orange circles) must all be present for browning to occur.

How are Arctic apples made?

Genetic material is stored in our DNA and divided into functional units called genes. The genes are “read” by copying the DNA sequence into a related molecule called RNA. The RNA copy functions as a blueprint that instructs the cell how to build the product for that gene, which is called a protein. The production of PPO enzymes, therefore, can be blocked by simply removing their RNA blueprints. To do so, researchers used a tool from molecular biology called RNA interference (RNAi). RNAi is a natural biological process that recognizes and destroys specific RNA structures. Biologists can use RNAi to lower PPO levels by introducing RNA sequences that cause the degradation of PPO RNA. Using this technique, researchers developed an anti-PPO gene that makes anti-PPO RNA, which destroys the PPO RNA before it can be used to make PPO enzymes.

Once scientists created the anti-PPO gene, they needed to safely introduce it into the apple genome. To make a variety called the Arctic Golden, researchers began with Golden Delicious apple buds and inserted an engineered piece of genetic material called a transgene that contained the anti-PPO gene. After confirming that the plant received the transgene, the saplings were then allowed to grow into mature trees, one of which produced the apple that is now known as the Arctic Golden.

Are Arctic apples safe?

After over a decade of research, regulatory agencies in the United States and Canada like the FDA and USDA recently approved Arctic apples for human consumption. Accumulated evidence shows that Arctic apple trees and fruit are no different from their traditional counterparts in terms of agricultural and nutritional characteristics. On the molecular level, the transgene genetic material present in Arctic apples is quickly degraded by your digestive system to the point where it’s indistinguishable from that found in traditional apples. The only new protein in Arctic apple trees—a protein called NPTII that’s used to confirm that the genetic engineering was successful—was not only undetectable in their apples, but it has also been evaluated and deemed nontoxic and non-allergenic by the FDA.

Yet some anti-GMO groups continue to protest the approval of Arctic apples, arguing that unforeseen consequences of the genetic alteration could impact safety. It’s true that it’s impossible to predict and disprove every possible consequence of a genetic change. But a recent review by the National Academies of Science that covers decades of published research found no convincing evidence that GE crops have negatively impacted human health or the environment. While it’s important to rigorously test all new crops that are developed, GE crops should not be considered inherently more dangerous than their traditionally-bred relatives.


So what’s next for the Arctic apple? It takes several years for new apple trees to grow and literally bear fruit, so it’ll take time for non-browning apples to expand to supermarkets throughout the US. Currently, Arctic Goldens are only available in bags of pre-sliced apples in select US cities, but Arctic versions of Granny Smith and Fuji apples have received USDA approval, and Arctic Galas are in development. If commercially successful, non-browning apples could help to combat rampant food waste one slice at a time.

Allison Baker is a second-year Ph.D. student in Biological and Biomedical Sciences at Harvard University.

For more information:

  • A cool time-lapse video that compares Arctic apples to conventional apples
  • This article on the history and economics of Okanagan
  • Multimedia resources that provide more insight into the science behind RNAi

Cover image credit: Okanagan Specialty Fruits Inc.

64 thoughts on “Arctic Apples: A fresh new take on genetic engineering

  1. Allison, you did a very nice job explaining the technology. It is so sad that there are probably thousands of genetic differences between the different varieties of apples on the grocery store shelf, yet people have become focused on one relatively trivial gene knock-out. Please keep up the good work; science needs all the good speakers it can get.

  2. I personally would not want to buy or consume the Arctic apple. Since genetic engineering is still so new, the long term effects on human health are still unknown and could be harmful. Humans have a right to be aware of the food they are consuming. Even if genetic engineering has no significant effects on consumer health in its current early stages, mass producing manufacturing companies will eventually take advantage of the increased profits of genetically engineered foods due to the decreased cost of production. Therefore, the freshness and nutritional value of fruits and other foods will continue to decline as a result of genetic engineering. An apple that does not brown due to its PPOs being removed is unnatural. Removing the PPOs from an apple is no different from removing the furniture from a house. A house does not need furniture, but it will no longer be comfortable or appealing without its furniture. To conclude, people will continue to waste food anyway, whether it browns or not. Most people do not save their food and prefer to eat it fresh anyway. Food waste is inevitable, but can be prevented through sustainability, recycling, and composting. Therefore, genetic engineering is unnecessary and will have little impact on improving the environment as a whole.

    1. Your poor biology teachers.
      Genetic engineering has been around for millennia in one form or another. Direct genetic modification is still decades old, and there is no reason to believe those methods are any riskier than any other methods.
      This includes nutritionally; contrary to the marketing you unquestioningly consumed, there is no nutritional difference between one of those eeeeevil GMOs and its “organic” counterpart.
      Your house analogy is deeply flawed. It’s meaningless, because taste doesn’t come from the enzymes. A more apt comparison would be a house without termites.
      And yes, yes fruit that doesn’t brown would reduce food waste. Sorry, but it is one of the major reasons fruit gets thrown away. It’s nice that you think everyone can compost their waste food, but leaving aside that absurdity – why is that better than not having to throw it away at all?

      Next time, actually try reading an article instead of reacting to it. Failing that, at least establish points using a logical sequence rather than a string of non sequiturs worthy of Natural News.

      1. Not the same as grafting and hybrids that are naturally bred to make a better resistant/stronger gene. These are DNA/RNA altered. Taking polyphenols out prevents them from allowing human and animal bodies from forming Quercetin that is anti-inflammatory, anti-cancer, anti-radicals. They do lose some nutritional value because of this genetic engineering. GMO is different and you are consuming RoundUP Ready herbicide even if in small amounts – if it is in everything you eat (processed foods of corn, soy, canola, etc.) can cause GI issues considering it causes an insects stomach to rupture from the inside out.

      2. I don’t know if you’ll see this, it being 4 years later and all, but thanks for your reply to her. It was very irritating to read her comment, but satisfying to read your reply. Couldn’t have put it better myself.

    2. While I don’t agree that GMO foods are any different from natural foods from the standpoint of health and nutrition, I think you have touched on two issues that are fair reasons to reject GMO out-of-hand, and that the burden of demonstrating that a GMO food is necessary rather than merely economically viable ought to be included in approving one for release into nature. First, genetic modification is proving to be an economic force that non-engineered foods cannot compete with. Canola, Corn, Beets, Soy, to name a few, are mostly GMO now. If you believe that economics is the only argument, then the discussion stops here. They win. When they are introduced, they will drive other crops out of the market. But behind that is the concentration of economic power in the hands of the patent holders. So long as their economic interests are dominant, then the utility of food crops, ie nutrition, becomes secondary to profit. That is dangerous, because it means famine may become secondary to wealth. The less food, the greater the price. and the greater the profit. But more importantly is the proliferation of the technology. A warning originated from the Asilomar conference in 1975 concerning this. There are two general problems. First, the more broadly this technology is spread, and the more variation there is, and this is especially true in these early days of CRISPR and gene drives, this will enter nature in a dangerous and unpredictable way. It takes little imagination to see what might result. Horizontal spread of toxin genes for example. And the studies that confirm that there has been no damage are necessarily looking at a few indicators of escape and damage, and also at what is being used now, and not at what will be invented. Each new technique and transgene is its own experiment launched on the world we all participate in. So these NAS assurances are quite empty. But moreover, there is a danger of the cumulative effects or synergy among transgenes. It simply isn’t predictable, but it is obviously dangerous. An A-bomb goes off once. Something like a run-away gene drive on newly fertile GMO ground goes off forever. And what is all of this for? So that we don’t waste apples because they look bad? So far what has happened is the appearance of pesticide resistant pests and GMO weeds. That is a tech solution for a psychological problem. Yes, we can feed the world by supercharging the green revolution with GMO. We could also feed the world without it. It would be better to change behavior, because I agree with you. Changing an apple won’t make people less wasteful. Quite frankly I think genetic engineering is full speed ahead for two reasons. Because we can, and because the iceberg hasn’t appeared yet. And money of course. And faith that nothing bad will happen. Faith based science. Another contradiction.

      1. Scientists don’t place their faith in a deity. They have faith in their own abilities. Believing in yourself doesn’t make one a hypocrite does it?

        1. Nice and informative post. I have subscribed to your posts and feed. As a marketing communications student, I must say that I found the post relevant to my subject area. Thanks, Helen

  3. Yes I’m sure that pesky enzyme causing browning doesn’t need to be there, 😂 God is laughing and thinks we are all so smart. 😑 Keep telling yourself Round up and all their nasty pesticides are fine that Monsanto now Bayer use to grow their toxic corn, soybeans and now apples. I’m sure it’s all healthy, 😉😉

    1. this isn’t about pesticides?? they’re not using pesticides to make the apples non-browning, it’s gene modification, so yeah it IS all healthy

    2. Hey do you have braces? Do you have glasses? Have you ever had surgery? You have a tattoo? Do you straighten or curl your hair? Do you put on makeup? God is laughing and thinks you are so smart. Why would God make you, you if you are going to change that. If god made you like that then why did you have to fix it. Doesn’t he not make mistakes, ok then don’t fix it because your sooooo smart.

  4. I am not a science experiment therefore I will not consume a science experiment. Not everything is better man-made. Nature knows better. We are turning our world into a fake toxic world. Science has no place modifying our food. Nature and organic, regenerative practices have mastered our food for a very long time now. We dont need big pharma and industry taking away societies last connection with the earth for more money in their pockets!

    1. then stop drinking water. science has “experimented” on that, purifying it so that we can drink safely. apples brown so that bacteria and fungi have a harder time getting a foothold on the apple. Since this apple isn’t going to be lying on a forest floor at the mercy of countless natural forces, all we did was take it away so that we would have less food waste. nature didn’t know that we would farm apples, reducing the need for this defensive mechanism. arctic apples were thought of and grown by an orchardist, not some giant pharma company or industry. -_- ?

  5. I don’t know much about apples, but I AM NOW VERY PASSIONATE ABOUT THEM AND IF WE DON’T STOP BROWNING, I CAN NOT GET MY THICC GRAMMY SMITH APPLES from the very sketchy “Farmers Market” across the street from me. SO STOP IT FASTER


  6. Fruit isn’t wasted because of some minor browning, its wasted because people are raised to be childish wasteful twits that take two bites and toss the rest. (evidence is that this applies to all foods not just fruit)
    I would also say that culling half the crop is just part of normal production and transport/storage not really “waste”, especially as fruit can be produce in far more abundance than people can eat and considering it is mostly just a source of carbohydrates. Protein is the main limiting factor in production of basic food staples.

  7. That’s because the Arctic apple is the product of complex new genetic engineering techniques that the USDA is just learning how to evaluate. … The Arctic apple has been engineered to silence the polyphenol oxidase (PPO) enzymes responsible for browning in apple flesh after the fruit is cut.

  8. With all this pesticide resistant GMO corn, amazing salmon that grows three times as fast and now apples that don’t turn brown, wow! To think that some hard working folks busted ass just for little ol’ me to have all of this wonderful food. And, they don’t even want credit for it! No bragging about their hard work advertised anywhere on their products. Be proud! What an achievement! Someone’s gone and “fixed” my food for me. Thanks so much! I didn’t even realize that it was broken.

  9. Excellent article! I would have liked to hear more about possible health benefits of eating non-browning foods. What are the chemicals produced during browning and how might they affect health? One example might be oxidized lipids, which are typically not a good thing. (free radicals). There was a nice review article in J Ag Food Chem in the 1996 by Friedman.

  10. Genetic engineering has been around for millennia in one form or another. Direct genetic modification is still decades old, and there is no reason to believe those methods are any riskier than any other methods.
    This includes nutritionally; contrary to the marketing you unquestioningly consumed, there is no nutritional difference between one of those eeeeevil GMOs and its “organic” counterpart.
    Your house analogy is deeply flawed. It’s meaningless, because taste doesn’t come from the enzymes. A more apt comparison would be a house without termites.
    And yes, yes fruit that doesn’t brown would reduce food waste. Sorry, but it is one of the major reasons fruit gets thrown away. It’s nice that you think everyone can compost their waste food, but leaving aside that absurdity – why is that better than not having to throw it away at all?

  11. Now we can eat rotten apples and not even realize there bad. This is the greatest thing since we started growing our own food

  12. I disagree with you because i don’t really trust GMOs yet since they are new but i do kind of think that GMOs are cool because they make your food bigger and that means there will be more food.

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