Triploid Apples: An adventure into their history, breeding and use

One of the most important considerations to me when growing apples in the South is if the cultivar has a tolerance to pests and diseases. Called “the final frontier” by my Northern and Western apple growing friends, the Mid-Atlantic and the rest of the US South are notoriously difficult areas to grow domesticated fruit. In true Southern hospitality, our soupy humidity and hot temperatures not only extend a warm embrace to all sorts of pest and disease here, but invite them to stay for a long while and breed.

Despite this high diversity of fungal, bacterial and insect pressure, there are still old apple trees in the landscape that have survived decades upon decades of environmental assault. These trees have been the subject and target of much interest in my network of fruit explorers, as these specimens are proof that it is possible to grow purposeful fruit and trees in this landscape without toxic, self-perpetuating inputs. In past essays, I’ve discussed rootstocks being a factor in this, where larger root systems tended to produce healthier trees.  But there are more factors in resilience than just the root system. In today’s essay, which has literally been in my drafts for 3 years, I want to discuss something I’ve been casually studying for years: Polyploidy, or having more than 2 paired sets of chromosomes.

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I’ll begin with a bit of history. In the early 1900s, there was a Swedish plant breeder and geneticist named Herman Nilsson-Ehle, who had spent much of his professorial career breeding wheat and oats for high yields in Sweden. He was a huge fan of Gregor Mendel, who had released his findings on inheritance only 8 years prior to Nilsson-Ehle’s birth, and his whole outlook on plant breeding research was a hat tip to Mendel. Mendel, for those of you who may be struggling to remember, was the Monk who stared at pea plants and developed the fundamental laws of inheritance, which we encountered in high school biology as the punnett square .

Before I go any further, I want to give a quick warning. From my research on Nilsson-Ehle, it appears he was a fan of “new Germany,” and saw the genetics research under Hitler’s regime as a means to save the world. In order to only showcase the apple breeding aspect of this man, I’m not going any further in this subject. If you want to read more on his thoughts, which scarily echo modern times, you can go here: Lundell 2016

In his early research of breeding cereal crops, Nilsson-Ehle would sometimes observe natural mutations in the hundreds of thousands of seeds he planted out for observation. These mutations had much larger, rounder leaves and after poking and prodding these mutants, he discovered their large size was due to having 2 additional sets of chromosomes, or polyploidy (Usually a diploid (2 sets of chromosomes), these plants were now tetraploid (4 sets of chromosomes). These plants exhibited giantism in all ways aside from vigor (which was relatively low). While the leaves and shoots were much thicker than diploids (2 chromosomal pairs), the flowers, fruits and seeds were nearly double in size. This was remarkable to Nilsson-Ehle and prompted him to theorize: If I take this mutant tetraploid and cross it back with its diploid self from the same cultivar, I should get a triploid (3 sets of chromosomes) that brings about enhanced genetics of both! 

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He was right. The tetraploids he crossed with diploids produced triploids that were more vigorous, hardy and resistant to disease than their diploid or tetraploid counterparts due to enhanced genetic modifiers inherited from the parents of two different ploidy (tetraploid and diploid). This brings me back to fruit exploring in the Mid-Atlantic and Southeastern US. The large majority of US cultivars known today as being able to tolerate fireblight, apple scab, powdery mildew, and loads of other issues while still persisting in the Southern landscape for decades upon decades are triploids! Including the Dula Beauty, my sturdy family apple cultivar.

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So the US picked up on Nilsson-Ehle’s breeding work and adopted it to their work in the states to breed for hardy, disease resistant apples, right? Nope. WW2 happened and we were already distracted with breeding for scab resistance (more about that in a bit). In 1950, famed berry breeder George Darrow reported on Nilsson-Ehle’s work in an address to the American Horticultural Society. In this address, he mentioned the premise behind Nilsson-Ehle’s work and connected the dots in how this way of thinking has translated into berry breeding for larger, higher quality cultivars. He briefly mentioned apples in this address, reporting that a tetraploid sport (mutant) of McIntosh had been found growing on branches of a normal McIntosh tree in New England, but the mutant branch was only half tetraploid, as the cortex of the wood was diploid (making it a ploidy chimera). He said they were trying to stabilize the McIntosh chimera as a full tetraploid through tissue culture, and I believe they achieved this due to the photo below. This was the end of an interest in sustainable fruit breeding in the US, in my grumpy opinion.

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Come on, Eliza, what about the Liberty apple? Goldrush? RedFree? Prima? [Slight rant/history on apple scab. Skip to below scabby apple pic to avoid]. Sure, there was a breeding effort between selected US land grant universities (PRI= Purdue, Rutgers, Univ. of Illinois) that began in 1926 to create scab resistant apples. They succeeded in doing so in a basic sort of way, which eventually led to the downfall of this research.  The style of their research was “monogenic,” or relying on a single gene to control scab resistance in an apple cultivar. There was also a whole lotta inbreeding going on.

The gene identified to have scab resistance is called the “vF gene,” which comes from the cultivar “Malus floribunda 821.” The reason why they picked this gene is because they could identify it in seedlings using molecular markers, so they didn’t have to waste time growing the trees to find out if it was scab susceptible or not.  That worked out well enough for a while and they selected some ho-hum cultivars (minus Goldrush, which is awesome but incredibly prone to cedar apple rust) to make available to the public. In 2002, the first reports of scab infection were reported on the scab-resistant apple cultivar ‘Prima.’

In 2011, a German pomologist wrote an article about all of this and, thankfully, it was translated into English shortly thereafter. What he found, looking into the lineage of most US and Euro scab resistant apple cultivars, was a huge amount of inbreeding going on. Not only that, but the cultivars being crossed back to themselves were highly susceptible to scab! I’ll quote directly from the article:

“Today the global fruit breeding industry is producing a wide range of varieties, with one big difference: the overwhelming majority are descendants of just six apple cultivars.

The author’s analysis of five hundred commercial varieties developed since 1920, mainly Central European and American types, shows that most are descended from Golden Delicious, Cox’s Orange Pippin, Jonathan, McIntosh, Red Delicious or James Grieve. This means they have at least one of these apples in their family tree, as a parent, grandparent or great-grandparent…” 

Many of the PRI releases have these 6 cultivars crossed multiple times in their lineage. If you do this right and bring out the right traits without problems, it’s called ‘line breeding’. If you end up with problems, it’s called ‘inbreeding’.

The second and main problem with this breeding work, in my opinion, was in our complacency with our selections. We basically ignored any further breeding efforts for scab resistance in order to pursue “Crisp” apples. Takeaway message: FEEL GUILTY ABOUT EATING A HONEYCRISP, COSMICCRISP, CRIMSONCRISP KARDASHIANCRISP ETC. BECAUSE THATS WHAT BREEDING LOOKS LIKE NOW INSTEAD OF BEING ABLE TO GROW APPLES WITHOUT MAJOR INPUTS! Too bad we haven’t been thinking about triploids or even multiple-gene scab control for the last 50 years.

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Guess who has? Russia. 

Since the early 80s, the All Russian Research Institute of Fruit Crop Breeding (VNIISPK) has continued with the scab resistant vF breeding work that spread across the US and Europe, only it is way more badass. Not only are they breeding for scab resistance, but they’re breeding for tolerance to late frosts, consistent yields without having to thin fruit, COLUMNAR growing habit AND Nilsson-Ehle’s version of triploidy (Speak a little more into my dirty ear, Russia). However, the near-sensationalism of these claims doesn’t stop there. Dr. Evgeny Sedov, the primary researcher in this endeavor (and someone I would really love to interview), closes the abstract of one of his scientific papers that goes into his triploidy research with the following that is so, so Russian:

“It is noted that triploid apple cultivars developed at VNIISPK are inferior to none of the foreign cultivars, based on a complex of commercial traits, and they significantly excel foreign cultivars in adaptability. Our apple cultivars may contribute to the import substitution of fruit production in Russia.”

Some mentioned and additional benefits of triploids (Or reasons to pursue more polyploidy breeding):

  • Adaptability to climate, disease, stress: In the above quote, Sedov writes how his triploid apple cultivars significantly kick other apple cultivar ass in terms of adaptability. And based on my research covering the last 100 years, he’s not wrong. There have been many observations by the scientific and lay community reporting that triploids end up being more cold hardy, more heat tolerant (the thickness of leaves and fewer, larger stomata give rise to a lower transpiration rate and more water retention that can be used during drought), have better nutrient uptake, and improved resistance to insects and pathogens. The theory for triploids having a higher environmental adaptability has to do with  an increased production of secondary metabolites, which enhance plant resistance and tolerance mechanisms (as well as chemical defense). 

  • Thinning: Triploids often have low fertility due to a reproductive barrier of having an extra set of chromosomes- making pollination difficult. Some apple pollen tends to pair decently well with triploid apples to get a decent crop. With most cultivars it isn’t great- just good. This could be seen as a boon to this class of ploidy, but I see it as a good thing. One of the greatest challenges to organic apple production is the thinning process. Most non-organic orchards thin using chemical sprays to knock off flowers or fruits. To this day, many organic spray chemicals either do a lackluster job, or oh-god-that’s-far-too-many-job of thinning the fruitlets off, leaving many orchardists to either thin by hand or accept biennalism (which was a 3 hour conversation at Stump Sprouts one year). If you have healthy pollinator populations, less fruit on the tree will guarantee you a return crop the next year, barring other environmental catastrophes (which you’re better prepared for with triploids, anyways).

  • Vigor: In the past, I’ve written about vigor on the Elizapples.com blog and how it’s my number one enemy in the Mid-Atlantic given my heavy soils, warm temperatures and ample water supply. Though I need to revisit those essays and condense them into my current evolution of thought, the reason for my past concerns around vigor is that I have conditions that induce [what I’d like to think is] “artificial vigor.” In my climate, this shows up as extreme vegetative growth, which sometimes gives rise to heightened fireblight pressure and other vulnerabilities. Though “artificial vigor” is likely what an incompatibility of growing conditions looks like, I’ve started to differentiate it from what I’m calling “true vigor,” or youthfulness through heterosis/hybrid vigor. This is where triploids shine.

    When you start digging in old texts, back before the rise of clonal rootstocks, you might encounter mention of two classes of trees referred to as “Standards” and “Fillers.” The “standards,” often mentioned as Baldwin and Rhode Island Greening (both triploids) were larger trees that took longer to bear fruit. These were thought to be permanent trees, or trees that would be around for generations. The “fillers,” such as Yellow Transparent and Wealthy, produce much smaller trees in the same length of time and were far more precocious in bearing fruit. These trees were thought to be temporary, and were planted in between the “standards” to increase production in the early life of the orchard. An unfortunate modern day “filler” would be HoneyCrisp (diploid). Growing in my climate, it is better termed runtycrisp. Super low vigor, gets loads of diseases, precocious bearer, dies early. Sort of an orchard mercenary. This, to me, is a good way to think about vigor. If you’re growing for the long-term, you’ll want a truly vigorous cultivar that teems with youthful energy, and I believe that youth is heightened as a triploid. If you are growing in areas that are full of pest and disease, it is also not a bad idea to have an extra set of chromosomes to help with defense and stress. Relic trees standing tall in the South tend to be triploid and their presence speaks to their youth and defense: Arkansas black. Fallawater. King David. Leathercoat. Roxbury Russet. Stayman Winesap. 

    With all of this said, we have a lot of work ahead of us to start thinking about what our breeding programs would look like if we set our targets on low-input, no spray, multi-gene disease tolerance and more. I get it, HoneyCrisp can store for a calendar year in my crisper drawer, but that’s all it has going for it after a year in there.

    I am pulling for the expansion of ‘process’ industries such as hard cider, vinegar, juices, syrups, etc to become the targets of agroforestry planning and planting enterprises in the near future. Annual or livestock farmers don’t want to mess with sprays or inputs that are outside of their normal non-tree crops care. If they are going to receive incentives to plant trees on their farms, they will want the ones that need little care and have an economic outlet. This will require a new set of apple cultivars to choose from and they have to come from somewhere…

     

Here is an incomplete list of confirmed triploid apples. Many of these are from the UK and do so-so in my climate. The ones with asterisks are what I have seen as old relic trees in the Mid-Atlantic:
Arkansas Black*
Ashmeads Kernel*
Baldwin*
Belle De Boskoop
Blenheim Orange
Bramley’s Seedling
Buckingham*
Bulmers Norman
Canadian Reinette
Catshead
Close
Crimson Bramley
Crimson King
Crispin
Dula Beauty*
Fallawater*
Fall Pippin*
Frösåker
Genete Moyle
Golden Reinette von Blenheim
Gravenstein*
Hausmuetterchen
Hurlbut
Husmodersäpple
Jonagold
King David*
King of Tompkins County
Lady Finger
Leathercoat*
Margille
Morgan Sweet*
Mutsu
Orleans Reinette
Paragon
Red Bietigheimer (Roter Stettiner)
Rhode Island Greening*
Ribston Pippin*
Roter Eiserapfel (Has 47 chromosomes rather than 51)
Rossvik
Roxbury Russett*
Shoëner Von Boskoop
Spigold
Stäfner Rosenapfel( Has 48 chromosomes)
Stark
Stayman*
Stayman Winesap*
Summer Rambo*
Suntan
Tom Putt
Transcendent Crab
Transparente Blanche
Vilberie
Vixin Crab
White Astrachan
Winterzitronenapfel
Winter Pearmain
Washington Strawberry

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How to Make Bradford/Callery Pear Less Invasive.

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In my last essay, In Defense of Bradford Pear, I showed the above chart from Australia that correlated fruit size with bird species. Similar charts or descriptions have been found in publications from New Zealand, Spain, Japan and in the US, as well. Based on the correlation of fruit consumption with fruit size, I’ve decided to elaborate on the last essay in order to practically address Callery/Bradford pear invasiveness in the US in the best way I can.

Cedar Waxwing eating Pyrus calleryana in winter. Photo from Pilot Online

Due to Callery’s fruit size attracting our native songbirds, like American robins, cedar waxwings and gray catbirds, we can’t stop them from eating the little pears and pooping in marginalized areas like fencelines and worn out pastures. To think we can kill enough Callery pear to make a difference is a lesson in futility because 1.) We live in the United States and you can’t go kill a neighbor’s tree in the name of INVASIVES if they don’t want you to and 2.) Each tree produces thousands of fruits. So, with that said, here are my top solutions to sustainably make Callery pear less invasive and more useful.

1.) Citizen Breeding. What makes Callery pear invasive is its ability to produce copious amounts of small fruits, which birds then eat and distribute all over the place. It seems logical, then, to want to try and breed larger fruits into our populations of Callery in order to stop the spread by birds. In order to reduce invasiveness by around 80%, all it takes is getting progeny from the Callery/Bradford trees to produce fruits that are around an inch (25mm) in diameter. How do we do that? Allow them to hybridize with larger fruiting pears so the seeds dispersed by birds will have a higher likelihood of growing larger fruits…thus halting the invasion cycle.

What is needed to hybridize these pears and get them larger? For starters, you’re going to need a collection of pears that bloom at the same time as Callery, which is quite early. Russian/Cold Climate and early Asian pears are likely your best bet for this, so I went through the GRIN database (taxpayer funded genetic repositories) and have made a starter-list (there are a bunch more):

PI 541904- Seuri Li
PI 45845- Yaguang Li
PI 437051- Jubilee (cold hardy)
PI 541925- Kor 2
PI 267863- Pingo Li
PI 134606- Tioma (cold hardy)
PI 278727- La Providence
PI 278731- Sivaganga Estate
PI 307497- Seu Ri
PI 292377- Ranniaia Mleevskaia (cold hardy)
PI 541760- Chieh li x Japanese Golden Russet
PI 278729- Samy’s Estate
PI 541761- Chieh Li x Japanese Golden Russet 2
PI 541905- Szumi
PI 127715- Krylov (cold hardy)
PI 541326- Angelica Di Saonara
PI 324028- B-52 (cold hardy)
PI 541290- Mag 1 (cold hardy)
PI 132103- Shu Li
PI 312509- Tse Li

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You can request free scions online from September 1 to February 1 of every year from GRIN. You can also probably buy many of these cultivars online. From there, I highly recommend you share scions of these for free every winter, as I plan to do, in order to help infuse larger fruiting genetics into Calleryana.

You might notice there are a bunch of Asian pears in that list and you might think: Eliza, those pears are super fireblight susceptible! And you are right, of course, but think of it this way: MANY trees that are listed as fireblight susceptible are actually quite tolerant to FB once they are established and reaching sexual maturity. With Callery being an amazingly fireblight tolerant rootstock, this should help to get your topworked trees past the first 2 years of heightened susceptibility so they can start to fruit. Once these Asian pears intermingle with Callery, there are two possible outcomes:

1.) The hybrid offspring are more fireblight tolerant than the grafted Asian pearent’s tolerance

2.) The hybrid offspring is less tolerant to fireblight than the grafted Asian parent’s tolerance and will probably succumb to the disease and die on its own.

Either are a win-win, really.

Next, you’re gonna need to go into your pear thicket and do some cutting and grafting. There are two scenarios I see often:

1.) Field full of Callery: If you have a thick field of calleryana, I would recommend getting a forestry mulcher in and cut/mulch rows into the existing Callery stand. Then, run the mulcher to cut out trees within the rows left standing so the remaining are at 15 foot spacings. Top the trees you’ve left behind above deer browse ( throw into the alley and run over those, too, with the mulcher) and graft on the early blooming large fruited cultivars.

2.) Fenceline/Border with Callery: This is the scenario We’ve been dealing with over the past few years along the farm fenceline. First thing I do is flag the trees I want to keep, which are at 15 foot spacings along the fence. Then we cut out and chip all the non-flagged callery trees using my neighbor’s chipper (I mulch my orchard with callery pear wood chips). While we are cutting out the non-flagged trees, I go ahead and also cut the tops out of the flagged trees. I pick a height that is above deer browse height and also has a lot of clear wood without branches, because that helps with grafting. In April (I’m in zone 7a), I make fresh cuts on the remaining pear trees and topwork all of them to fruiting cultivars. We’ve been doing this for 3 years and 2018’s topworked pears will be producing fruit this year.

Topworked fenceline callery pear to a local french heirloom cultivar. This was grafted in April of 2021
Topworked fenceline callery pear to a local french heirloom cultivar. This was grafted in April of 2021. This is a smaller tree. I’ve topworked 7″ trees as well with amazing take.

This is totally doable and the result? An orchard of pears! You’d have to cut the tree down anyway if you were going to spray it, so why not turn it into a producing pear tree of value? My neighbors even pitched in to help us cut and chip in the name of supporting my vision and also getting rid of the fruiting portion of the Callery trees.

In two years, your top-worked pears will be flowering and the bees will mingle between surrounding landscape Callery/Bradford pears that weren’t able to be cut down and the large-fruited cultivars you have grafted. With callery pears being pollinated with the list of pears above, your chances of getting larger fruit to come up from the fertilized seed will exponentially increase, limiting its invasiveness if the fruit is an inch or larger in diameter.

2.) Use them as rootstocks! Every Callery pear growing is automatically the best pear rootstock around. For all of you people out there who are inundated with deer pressure, graft to the Callery pears to any pear you’d like (or Winter Banana apple) above the deer browse line. Sure, you’ll get lots of leafy re-growth off the trunk for a few years (which the deer or other livestock eat as tender shoots), but its also really easy to remove new growth with your hands or slightly older growth with pruners, and new shoots don’t have thorns. You’ll start to get fruit in 2-3 years.

One of the main reasons why Callery didn’t catch on as a rootstock, aside from root propagation failures and hardiness, is that they don’t produce dessert fruit (fruit meant for out of hand eating). This is the same reason why we’ve lost SO MANY fruit cultivars in the last 100 years. If you weren’t a dessert cultivar chosen by the cooperative extension to be grown in the early 20th century, you were phased out. However, in today’s markets, I believe large fruited Callery pear hybrids really have a chance in fermentation, specifically cider blends and perry (cider made from pears). They are high in sugar (over 16% brix on average for the 200 or so hybridized trees I’ve evaluated), and run the gamut in acidity, tannins, aromatics and unusual characteristics. Since these trees are so disease and pest tolerant, which allows them to grow and produce copious amounts of fruit without the hand of humans or chemicals, they stand to produce the most sustainable fruits and alcohol in humid temperate climates. We need more people working with them in order to make this happen because they aren’t apples and they need their own methods.

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The last essay left me with a bunch of hate mail and loads of baseless claims. In future essays, I’ll be debunking many of these claims in order to try and bring about a full picture. With that said, please send your strong opinions to fruitandfodder@gmail.com

Of Note: throughout the South and Southern New England, I have been noticing spontaneous hybridization in the “wild” between P. calleryana with P. communis (French) and/or P. pyrifolia (Asian). These trees have much larger fruits, usually golfball sized or larger, and are often loaded with fruits dripping from the trees because Callery genetics are heavy lateral bearers (perhaps an indicator phenotype for these hybrids). No research that I can find has evaluated the genetics of these larger fruited callery-like pears to see what exactly they are crossed with, but I’m happy to help supply specimens if anyone out there takes an interest.