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* (struggles with brown rot)
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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Cider And Heirloom Apple Vigor: An Hypothesis

Recently, I was on the phone with a mentor and we were discussing hedgerows (my new pet project, aside from brewing all sorts of alcohol). With some of the species I mentioned, I was told that livestock would eat them down to nothing and render the hedgerow useless. After having a few tree species rejected, I frustratingly asked: “What if I planted my hedgerows with invasives like multi-flora rose, then?!”

Without any hesitation, my mentor said: “Invasives like multi-flora rose are very delicious to many animals, like my goats.  You might be suggesting invasive plants for your hedgerow because they are vigorous and seem to outcompete everything else, but try to think about vigor from another perspective. If plants with high vigor are also the most sought after by animals, don’t you think that vigor might be an evolutionary trait to survive browse?”

This is the first time I’ve heard this perspective on invasives and I’ve really enjoyed thinking about it. After some minimal research, I found out that the subject is still debated today by ecologists as the “plant vigor hypothesis.” Generally speaking, vigorous plants have higher nutrient densities than non-vigorous plants, so herbivores are more prone to eat them. However! If the very vigorous cultivars are able to put on a bunch of girth, many herbivores aren’t able to eat the whole thing because of their jaw size.

This, of course, has got me thinking about apples. Here’s why.

In many essays on this blog, I’ve talked about how I consider many cider and heirloom cultivars to be very vigorous as compared to most of the grocery store cultivars. Vigorous cultivars are harder to prune, occupy more space (so less trees per acre), have issues with vegetative vs fruit bud proportions, etc. In general, they are harder to grow.  After reading more about this “plant vigor hypothesis,” I wonder if there is a connection between vigor and nutrient density in apples cultivars?

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From an evolutionary standpoint, a correlation between vigor and nutrient density makes sense to me. Many wild crab apples in the US have much higher tannins (aka polyphenols, which =nutrition density) than cultivated varieties. This is from the many lifetimes spent co-evolving with insects and herbivores who are trying to eat them. From observing crabapples in the “wild” and planted in landscapes, it seems as if many trees have low vigor and perhaps this is because they have evolved to have an unpalatable deterrence for animals and humans alike?

In hard cider, many of the wild crabs are too much for our palates to handle and though very nutritious, they will cause a harsh and likely negative consumer experience. So what have we done? Over time, cider drinkers/makers/apple growers have selected cultivars to grow which are palatable to the consumer, but also contain enough tannins (or polyphenols, or natural defense) to give the cider some substance.  Could it be that in selecting not-so-astringent apple cultivars for eating/drinking, we’re unknowingly selecting for more tree vigor? If the apple cultivar hasn’t evolved enough to deter herbivores through astringent taste, then do genetics dictate that it must rely on vigor to survive? 

These sorts of questions make me excited and I’ll keep learning about these processes in order to try and uncover different management ideas that don’t involve regulating vigor through the use of dwarfing rootstocks, black magic hormonal potions like Apogee (which converts vegetative buds into fruiting buds), and planting in light soils. All of those management aspects, I suspect, are making the vigorous cultivars less vigorous/more fibrous/less nutrient dense.

Thoughts to be continued, but in the meantime here are a few off the top of my head:

Thought 1: Pruning extremely vigorous varieties like an herbivore in order to get faster fruit set?

Thought 2: Continuing to fruit explore to find mixes of wild x cultivated which hit high nutrient densities, palatability, and lower vigor.  (I’m writing a fruit exploring book about how to do this at the moment)

Thought 3: Making crabapples a significant part of my home breeding program.

 

 

Fruit Exploring: Hunting the Vermont Beauty

Today I ate 6 pears, 11 apples. I tasted 14 pears, 24 apples. I managed to drop off apples at my rendezvous point on time for the delivery guy, and all else after that meeting at noon went down hill. Tis the season to be perpetually late to appointments/meetings due to the abundance of fruit on the roadside.  Also: Tis the season for elevated fiber consumption and trying to deal with it.

I’ve wanted to write a blog post about fruit exploring for ages now, as it is an exciting and integral part of my fruit life. Today’s essay will only be a brief glimpse, as I am writing a book on the subject at the moment. But I wanted to take a moment to tell you about my Tuesday.

Earlier this year, I was putting away the “Pears of New York” book by U.P Hedrick and had this impulse to open it and look at one of the colored plates. When I opened it, the pear I landed on was called “Vermont Beauty.” I’m right across the lake from Vermont and one of the orchards I work with is in Vermont, so I decided to read on…

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From this book, I learned that this pear was a seedling planted out by Benjamin Macomber of Grand Isle, VT…an island in Lake Champlain. I currently live across the street from Lake Champlain! So, I got to hunting. The first step in fruit exploring is research, which occupied an entire Sunday. After having an idea of where to look for this pear, I called my good Vermont friend Meg Giroux and we went fruit exploring for this pear over Memorial Day Weekend.

Fruit exploring involves a lot of cold-knocking. “Hi. My name is Eliza; I’m a fruit preservationist and I have a story to tell you [insert story for specific fruit]…. Do you know if you have any fruit trees on your property?” That is the usual gist of my approach, full of smiling and excitement. Only once, in Georgia, have I been turned away…

We were looking for the remnants of an old 40-acre orchard we discovered through a local Georgia historian who knew a lot about peaches, brandy and a particular Southern family’s plantation drama/legacy. Thanks to his directions, we were right in the epicenter of where the orchard once was and we decided to go door to door knocking, seeking permission to check out the apple trees possibly growing on their property. We knocked on one door and the people were obviously home, just not answering. I left a message on their car about why we were there, how we’d love to look at their orchard (which is the only remnant we could find of the once 40 acre orchard), and my contact information. I got a call later that day: “Hai, I’m lookin’ for Elyyyzuh. My husban’ tol me to call yew and tell yew that yew are not welcome on our properteee.” After I tried to tell her that we were only interested in their apple trees, she kept interpreting/spitting condemning words for her husband, who I could hear in the background.

Upset and saddened by their isolationist/paranoid ways, I wrote the pastor of their church (because we previously talked with the home owner’s mother who told us they belonged to the church on the hill) and asked the pastor to preach a lesson on “Do not neglect to show hospitality to strangers, for thereby some have entertained angels unawares.” (Hebrews 13:2). And then proceeded to tattle on the couple, because if anyone knows me well, you know that I have band of “angels unawares” by my side.

Anyways, back to Vermont. After knocking on every single door we thought might have a property with fruit trees attached to it, we were given wide-spread permission to look around. We found an old, old pear. An old old apple. Another old property with an old pear and apple, etc. And after finding a map from the 1800s, we found an old, old apple orchard. Excited, I promised myself to go back in the fall when they had fruit. AND BOY DO THEY HAVE FRUIT.

This story will be about the first pear tree we found, which I checked first with fingers crossed in hopes that the fruit wasn’t an early season ripener. Luckily, it wasn’t and the tree was loaded with fruit which had just decided to start dropping (!).

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I had every hope back on Memorial Day that this might be the Vermont Beauty. And, well, it may be. An extension agent recently told me that cooler temperatures instigate better color in fruit. With the downright hot fall we’ve had up until a few days ago, that could be a reason for the lack of color on this pear. Otherwise, it seems to match up decently well with the description found in “Pears of New York.” The flavor is sweet, very fine grained, has potential to be buttery-melty in the mouth once it has been given a chance to fully ripen off the tree.  I might be overly hopeful, but…

I stopped at three other properties on Tuesday, each one tied for having the oldest pear trees I’ve ever seen. Here’s one of the trunks (shoe for scale, it was completely hollowed out and probably close to 4 feet in diameter)…

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And you know what? ALL OF THE PEARS ARE THE SAME KIND!!!  They all taste the same (minus some minor soil differences); They all have that light blush on them; They all have the same shape.

I’ve since done some background research of these properties and they line up with my research as having a connection with Benjamin Macomber, the cultivator of the “Vermont Beauty.” I can’t reveal this connection just yet because it’s a chapter in my book, but just trust me for the time being that they are all connected. The rest is left to old books with descriptions so I can try to key this pear out. The trouble is finding an adequate description of it!  If in fact this is the “Vermont Beauty,” I have just found a lost variety. In order to save it, no matter what it is, all property owners have given me permission to take cuttings this winter from their trees. We might never know the name, but it has proven itself to be resilient for at least 100 years (I think closer to 200), and that is worth saving and propagating.

I hope to graft this tree this year. Some of you might ask for scionwood, which is quite alright with me, but my team and I have been trying to come up with a way to both spread the goods and retain some funding for our future fruit exploring missions.  With that said, the proceeds of this scionwood and future tree sales will go into #thefruitexplorers research fund. We poor-yet-passionate heirloom/resilient fruit nerds will ask that every propagation of this pear (or anything else we find) will spur an on-your-honor royalty donation on our soon-to-be website…and that it’s story be passed on to the new owner (who, if they decide to propagate, will carry on with the royalty donation).

Stay tuned, we have found some amazing specimens this year and there are more to come (we have loads of self-funded research that, of course, costs money). Already, we’ve found a Southern bittersweet crabapple out of Alabama which I think could be the next great Southern cider apple, showing awesome pest and disease resistance. Pending is to see if it has an annual bearing tendency.  Another is a gorgeous, insect and disease resistant (from what I can tell) “sweet” apple that lacks acidity, which Benjamin Watson touts as a great ingredient to blend with high acidity/sugar American heirlooms. Many more…we just need to get our acts together, which is a winter task.

Thoughts? Would love to hear.

basic information young farmers should know about me

My name is Eliza Greenman and I’m an heirloom and cider apple farmer in the Blue Ridge Mountains of Southwestern Virginia. I believe in opening up about the following information and I wish more farmers would do the same. Let’s get rid of the smoke and mirrors, eh?

  • Age: 30 (until December)
  • Businesses: I own an apple tree nursery business called Legacy Fruit Trees (it’s my first year). Other than that, my future orchards are still in my nursery or have yet to be grafted.
  • Do I come from a farming family? No. I come from the [now] suburbs of Southeastern Virginia. My Grandparents once lived on a farm, but they were forced to sell it to the city because of its prime location. They also didn’t farm for a living.
  • Do I come from “family money?” For US standards, I would consider my upbringing to be that of middle class.  No trust funds, no direct inheritance, no secret piles of money that I know of.
  • Do I have a family of my own?  No kids. No husband. No boyfriend right now (I’ve been too busy to date). I had a dog but that’s a story for another post.
  • Do I own land? Nope, sure don’t.  I’m involved in a series of creative arrangements that basically work out to being a long-term lease. I will talk about this in full later. It’s a long story.
  • Do I have an off-farm job? Technically, no. I live and work on a farm that is home to a hard cider company. I manage their hard cider orchards 2 days a week, which amounts to around $850/month. Lots more to come regarding that relationship.
  • Did I go to college?   Yes, I went to a 4 year university and earned a BS degree in Forestry.
  • Did I graduate with student loans? I graduated with $20,000 in student loans and managed to pay them off last year (2013).  $10,000 of those loan repayments came from Americorp’s Educational Award.

Ok, I think that’s a start in transparency. Any questions, don’t hesitate to ask. I solemnly swear to be as honest as possible in this blog.