Wednesday, October 22, 2014

Some Thoughts on My Program...

Some Thoughts on My Program and Objectives

Breeding With an Eye to Many Traits

As this year is drawing to a close, I have been reviewing my program and my objectives. I find I am very pleased with where my program is heading. I see a lot of potential even though a great many of my seedlings are just bridge plants to be used in further breeding work. However, amongst those seedlings, I see many of my goals beginning to come together and I am encouraged that the images I see in my mind's eye can become a reality.

It is becoming apparent to me that because I often write about breeding for disease resistance with a focus on daylily rust, there are a good number of people out there who seem to associate my program strictly with rust resistance breeding, but that is far from the case. While resistance to rust is important to me, I simply view that resistance as one trait in a larger set of plant traits I consider important and am selecting for. However, just like every other daylily breeder on earth, my most exciting end-goals revolve around the flowers. Let's face it, we aren't growing daylilies as strictly foliage plants (even though we are stuck with just their foliage for most of the year - a thing to consider).

With that said though, there are many plant traits that I find equally important to the flower, and a beautiful flower on a terrible plant is a very disappoint thing for me. So my goals are beautiful flowers on attractive, hardy, well-performing plants. I see that as a complete package. Now I will stress that I am realistic. I don't believe there is such a thing as a completely perfect plant that has every positive trait you could ever want and will be that perfect in every environment and garden it is grown in. Sure, the attempt to produce such plants is a worthy goal and the Lenington award recognizes the best of such plants, but I doubt there is any one cultivar that is equally perfect in every situation. If nothing else, varying tastes in flower types will potentially discredit some of the best for some people.

I have a broad taste in flower types. I love spiders on the one hand, and ruffled full-forms on the other, unusual forms on the one hand and sculptural types on the other. Teeth are amazing and patterns are stunning. Eyes and edges entice and yet a good, bright yellow or near white self is a delight in the garden with carrying power and marvelous grouping effect in the landscape. I see a use and need for most of the phenotypes that exist in the daylily world and I am working with a wide range of phenotypes. I am not in any particular camp as to the 'right' flower type. I can't choose, I love most of them and I see different uses for them all, and for that matter, I like both diploids and tetraploids.

However, from growing daylilies for nearly forty years, I do have a long list of things I don't care for. Plants that don't increase, poor bud counts, low scape to fan ratios, weak scapes that lean or fall completely, ugly foliage, flowers that don't open, flowers on scapes so short that they are down in the foliage, muddy 'not-really-any-color' colors, canoed petals, spotty petals, high disease susceptibility, and infertility, are examples of traits I greatly dislike. I find all of these things disappointing, so in my own breeding program, I make every effort to cull out these undesirable traits when I find them in my seedling beds or the hybridizing garden.

Yet, there may be instances where any given seedling or cultivar is so good in one or two traits,  that I can overlook its other lesser traits, and I think we all have such instances. Such plants are regularly used in breeding programs, but it is nice if we can make the attempt to combine the good traits we so value with other good traits that such a plant may lack in an effort to make superior plants with many desirable traits. That is called breeding. So the goal of breeding is generally to produce superior plants with more and more advanced flowers, but ideals cannot always be realized in reality. That doesn't mean we abandon the attempt, though.

What it does mean is that for anyone who is being realistic and hopes to ever introduce a cultivar in their lifetime, there are always compromises, as the notion of the perfect plant may be more of an ideal than a reality. While it may be an ideal to always be reaching for, compromise may mean that you introduce something that is marginally less than perfect, or even if it is perfect to your eye or in your garden, you recognize that it may not be so to other people's eye or in their gardens. These things have to be accepted when we are dealing with an organism that is as environmentally influenced as the dayliliy.

Very few of us have the resources to breed plants as the professional plant breeders do or to utilize all of their techniques. We may incorporate what we can from their techniques and make our best effort, but we also have to recognize as hobbyist breeders that we all have some limitations. It is important to assess our abilities honestly, so as not to build unrealistic expectations that may well lead to disappointment - either for ourselves or others. We are probably all a bit unrealistic when we are getting started, though time and experience usually help to adjust that earlier idealism.

As an example, the beginner may say they only want to produce plants with six or more branches, or thirty or more buds, but then you get an amazing flower with four branches and fifteen buds, and you just can't cull it, so there is a compromise. We might make excuses, but the best approach is just to be honest - this flower was too pretty to cull. Now if you want to stick to your guns somewhat you might make that plant a bridge plant, breeding it to find an offspring just as lovely but with better branching and bud count. However, some may choose that plant as an introduction while others might use it is a bridge plant only and neither are really wrong. It is there choice and the marketplace will determine whether the breeder was right or not.

Some traits may have more importance, but I wanted to start with a flower example, because that is where most of us leave our guidelines or rules behind. However, it is good if we can be less easily seduced by pretty faces. When a plant has a serious flaw (and the above example is not really a serious flaw) it is probably best if we can look past the pretty face to the flaw and either discard the plant or only use it as a bridge plant, carefully testing its offspring to find those with its good traits and without the major flaw. I suspect if this had been done more, there would be less disappointment regarding some cultivars. One thing I am finding though is that there are many pretty faces produced in every seedling crop, so personally, I don't find it impossible to cull a pretty flower if there are other serious flaws.

My philosophy is that I let the plants guide me. I set out a group of criteria as a guideline in my breeding and I eliminate those that don't fit the criteria, but I do try to make that set of criteria relatively realistic and I do not expect every plant to match all the criteria points. Some of the selection points are more important to me than others. However, beyond that static set of criteria, I observe what is actually occurring and let that be my main guide. Whatever is working is a gift and should be exploited, whether it fits our narrow expectations or not.

This may be a good point to discuss my actual goals with rust resistance. Classical disease resistance selection would focus solely on producing the most resistant plants, but I look at my program as being focused more on actively removing the most susceptible plants from the gene pool. I certainly note the recurrently most resistant cultivars and seedlings, and I note this every year I have rust, using those that show the most resistance each year as major breeders, but my goal is to produce reasonable garden resistance rather than the most extreme immunity that is 'forever and always'.

I consider my goals and methods to be more realistic because I can't use all aspects of a professional-level selection system as would be used in commercial operations and rust is constantly mutating so that a highly resistant/seemingly immune plant in my garden may not be equally resistant in another garden with another strain of rust, but for that matter a fast-increaser in my garden might be a slow-increaser in another garden or a plant with strong scapes in my garden may fall in another garden or on an immature division. I just find it to be important to be realistic as I pursue my goals.

All I can ever say is that a given cultivar I might introduce shows a given level of resistance in my garden for 'x' number of seasons. I can report the feedback I receive from others, and all of this is relevant information, but it doesn't mean that any given cultivar in my garden will perform the same in another garden, so it seems to me that the pursuit of rust resistance, and nothing else, is not a good use of my time and effort. I will say that I suspect there will be some cultivars that will show high levels of resistance in the face of many strains of rust and may exhibit that resistance for many years, but none are likely to be resistant in the face of every possible strain of rust or for all time. That is reality and to ignore it is to set myself and others up for disappointment. Again, I choose to be realistic.

Now, with that said, I still think it is important for me to select against rust susceptibility, as high disease susceptibility is a trait I find very undesirable, just as I find falling scapes, muddy colors, or flowers that won't open to be highly undesirable traits. So all these traits fall into the realm of 'things I cull against'. By removing such traits, I am working to move my gene pool in the opposite direction - toward disease resistance, strong scapes, clear colors, and flowers that consistently open well, etc. I see 'rust resistance' as no more or less important than any other trait. It is just another trait within the overall selection criteria I have set for myself and it is not, by a long shot, my only focus.

With the publication of the paper Identification of Pathotypes in the Daylily Rust Pathogen Puccinia hemerocallidis by Dr. Buck of the University of Georgia, we now have strong evidence that there are multiple strains of rust in the US. While this is no surprise to me, the publication of this paper gives us evidence of this obvious point - rust continues to mutate. The most encouraging point of this paper though is that there were a handful of cultivars tested in this study that showed good to high resistance to the isolates (potential genetical strains) of rust used in the test project. This is very encouraging and indicates that cultivars of broad-based genetic resistance do exist and can likely be bred. (I will likely offer more detailed look at this research paper in a later blog post.)

I believe that we in hobbyist breeding settings can make an impact through identifying cultivars with broad-based resistance and breeding value and any interested party should use whatever testing methodologies they feel comfortable with to work toward identifying such plants to breed from. As introductions are made from such plants I do feel it is important to report on the resistance that has been observed, without making that a major marketing strategy.

It may seem counterintuitive, but I don't really see rust as a 'premium' trait, in and of itself. While we may in time realize that some cultivars and the genes they carry offer broad-based, genetic resistance for a long period of time, there is probably no real way to know that or make such a claim as only time will tell. It is only in hindsight that we will know this to have been the case. Working with those that have been consistently resistant or seemingly-immune for a long period of time and in many locations does give us the best bet for long-term and broad-based genetic resistance to many strains of rust though. 

I see rust resistance as an issue of cosmetics. I feel this way because rust is generally not deadly (except for some instances in the deep south) and it only creates an unsightly mess. While this may be uncomfortable to look at and completely unacceptable for many gardeners, it is not a plague. So I tend to view rust susceptibility as merely another undesirable trait to select against, much like any other undesirable trait we would select against, such as spotting of the blooms or falling scapes (both cosmetic/aesthetic considerations).

In breeding for any trait, I find it very important to test potential breeders, noting their combining ability and breeding value, so as to locate those breeders with the best chances of passing on desirable traits and breaks in phenotype, while not passing on undesirable traits. This would apply equally to all traits. I don't see one trait as paramount and of more importance than any other trait. I am seeking balance and a balanced approach. Imbalanced approaches tend to allow a lot of small problems to accumulate over time through being ignored, while all focus is going to one or two traits. In time those many little problems can add up to major problems. That which is ignored does not go away.

So in my own program I am not seeking to go to any particular extremes with one single trait, but to create balanced plants with beautiful flowers. So while I do watch rust resistance and go out of my way to select against rust susceptibility that is not, nor can it be, my only focus. As an example, I am just as likely to cull a highly resistant plant with horrible flowers or some other serious flaw as I am to cull a high susceptible plant with beautiful flowers. Selection is a sword that cuts both ways. Or, I could say that I don't suffer serious flaws gladly, whatever those flaws might be.

In my program, I produce thousands of seed every year. In the first years, I produced around 20-30,000. Last year I produced 70,000+/- and this year closer to 100,000+/-. Yes...that is ridiculous. However, there is method to the madness, as I am selecting for many traits and only keep a few seedlings from any batch for any length of time. So it actually makes sense for me to have a long list of criteria for culling. The more traits I am selecting for, the more plants I can potentially cull and that allows me to get down to manageable numbers more quickly and potential be producing really superior plants in the process. 

I actually don't like to breed from huge numbers of individual plants, preferring to focus my efforts on a few plants that are well tested, but to get there, many plants have to be tested and evaluated. In time, of course, I will produce fewer seeds from fewer but more well understood cultivars of high breeding value. Until then though, I actually need a lot of things to cull for, just to reduce the numbers of seedlings I have, and of course, this also allows there to be a few seedlings with a high concentration of many desirable traits and/or real breaks in the flower on superior plants. That is the greater goal anyhow. So it makes sense for me to use rust resistance/susceptibility as a culling and selection tool. I can always stand to cull out a few more seedlings!

Culling starts early, continues as long as the plant is here and is not just focused on the flowers, as I cull lots of seedlings for various reasons before I ever see the first flower. However, I do not turn over my plants every year or two as is common in many operations. I tend to get first flower in two years, with some taking to the third year and only a few in the first year. (Those which take more than three years to produce the first flower are simply culled as well.) However, I find the flowers from the first year of flowering to be a highly unreliable guide to cull by. I have seen too many examples already of a flower that is gorgeous in the first year if flower only to look horrid the next to believe that selection based solely on the flowers of the first year of flowering is a wise idea. What I look for is a plant that has lovely flowers year after year. To do that, I need to evaluate the plant for several years.

Five years is my average evaluation time, both for a new cultivar I am considering for a major position in the breeding program and for my seedlings. My oldest group of seedlings, mainly from seeds I purchased in 2010 will be five years old next fall. I have been culling on those seedlings now for four full years. They have been through rust culling and foliage culling and flower culling. Of the 7,000 seedlings I started with, there are now about 100 plants left. Of those, there are four or five that I am considering for registration in 2015 and introduction in 2016 and maybe ten or twenty more that are likely to be retained as long-term breeding stock. I can't say I am completely sure yet on any of them though.

The first round of seedlings I produced here were from 2011 breeding and are now three years old (they were planted the summer of the year they were produced and were small seedlings before winter). They have been through several rounds of culling, but flower culling only began on these in earnest this year and will predominantly be done next year. Once they reach their fourth and fifth years, I will have a much clearer idea of their destinies, and of course there will be far fewer of them by that time as well. I am not sure any of these will make the cut for introduction, as most were created specifically to be bridge plants, but time will tell. I have now bred from a few of them in the 2013 and 2014 seasons and am beginning to gather preliminary data on the breeding value of some of the more promising. This is part of the five year evaluation and is another point of consideration - breeding value. This will determine who continues on in my breeding program and who gets composted.

While breeders further south are able to produce an introducible number of mature fans very quickly, and norther breeders can often do the same with greenhouses, I don't feel anyone can really understand their plants without a few years of observation, especially those of us growing in more temperate climates and strictly growing outside. Yet even if I were in the position to introduce plants more quickly, I think I would want to take my time and be as sure as possible about the consistency of any given plant and its flowers before I made an introduction. I think this would help me to avoid unpleasant surprises on down the line, though no one can foresee all possibilities or make perfect decisions all the time. We will all make mistakes and fail at times. These are gifts that we can learn from and no daylily is going to be perfect for all time and in every location, no matter what efforts we make toward that end. This may be one of the hardest lessons of all to learn.

In closing, I hope my thoughts have offered you insights into my process and perhaps also given you food for thought that may help your own programs. In some ways, many of my posts are more missives to myself, sort of a way to refocus or sort out my thoughts, than they are directions for others. They are sort of 'me talking to me' while allowing you to see the process. I do hope it can be of some help and guidance to us all :-)

Thursday, October 16, 2014

Lessons In Persistence

Lessons In Persistence:
Breeding for Rust Resistance

An Interview With Ron Reimer
Part 1

 Ron in his garden in Russellville, Arkansas

Since Ron has just this fall made his first series of introductions for 2015, I am rerunning the two-part series on his breeding program that I published in 2013. To see Ron's new introductions, CLICK HERE. To go to part 2 of this series, CLICK HERE.

Ron has also asks me to add a bit of new information to the beginning of this post concerning his current rust resistance ratings and views on resistance.

"Our past several years have given us extremely bad rust infestations.  This year was by far the worst due to repeated small rains and high humidity with heavy dew throughout the season.  This is unusual for us as we usually bake in high heat and drought for the summer.  This year's weather conditions were so conducive to rust infestation that it provided a super-acid test for rust resistance.  Lots of downgrading in the rust ratings had to be done, I'm afraid, and many outstanding cultivars got a blue tag, which is ultimately a ticket to the compost heap.  This too is part of the program, keeping a small population of rust prone plants handy in the garden, but not too many. On the bright side, many cultivars have remained free of rust throughout the season."

"The University of Georgia discovery of many strains of rust have prompted me to qualify my labels of rust resistant and high rust resistance.  On my photo gallery I have indicated that I cannot claim total immunity to rust; it is just more complicated now.   I still do have totally rust free daylilies in a rust infested garden, but I am less certain about their immunity in the face of uncertain climate and the multiple rust strain question." 

Ron's current rust rating system 10/20/2014

This is the first of a two part series on the breeding work of Ron Reimer of Russellville, Arkansas. In this first part, we will look at Ron’s breeding efforts toward producing a line of rust resistant diploid daylilies and in the second part, we will look at the products of his work thus far and the direction he is pursuing in developing a line of resistant daylilies of distinct phenotype for future introduction to the public.

In my pursuit of information on breeding for rust resistance in daylilies, I have found little information that is very useful and what information that is out there that is useful has been hard to find and has required a considerable amount of sleuthing to uncover. In general, when one is seeking to breed for a given trait, you turn to both breeders and researchers to gain knowledge of practical selection methods and academic and industry research. However, in the daylily world and the academic and industry setting, very little has been done, either in terms of practical breeding efforts or applicable research. So it was that I was very happy to begin to read posts on the daylily main Internet robin by such people as Matthew Kaskel and Ron Reimer. Matthew had pursued breeding for rust resistance in tetraploids, while Ron Reimer has pursued the same in diploids. I soon learned that Mr. Kaskel is no longer breeding daylilies, so that left only Mr. Reimer for me to communicate with about an active program.

I contacted Ron Reimer in late 2011 and began to actively communicate and over that time, I have been lucky to be able to ask him many questions about his program. What follows is information that has been collected from some of his robin posts as well as through our many conversations, when I have been able to pose many questions on his program. As I have over twenty years experience in breeding for resistance to various diseases in animals, I was able to pose many questions that gave me good insight into his program and helped to confirm to me that Ron has indeed developed rust resistance to a high level in his project lines.

To begin with, let’s look at how Ron got started with this project and a bit on his background with plants and plant breeding. In his own words, Ron says, “My horticultural background goes back to when I was about eight, visiting with local breeders and nurseries in the San Francisco Bay area.  I started to raise orchids when I was twelve and only stopped when I left Santa Barbara at age 35.  My degree is in Horticulture and Landscape Architecture.  I designed and installed landscapes for most of my life. Plant breeding has always been my first love…For many years, orchids and later here in Arkansas, apricots, crape myrtles, daffodils, and azaleas and last, but not least, daylilies.  I started hybridizing daylilies in 1989 with the idea of producing strong superior garden-performing plants. I had been working with both Dips and Tets, but preferred Dips.  My goals were to develop very large, wide-segmented, heavy-textured, flat-opening and flat-formed flowers.  I pursued early morning openers with clear very strong colors (like impatiens), which were free from Spring Sickness and root rot.  All colors of flowers must perform in our hellish summer climate and still look good at 8:00 p.m.

“I was close to introducing several Tets from my own converted seedlings when Rust appeared.  It was very devastating to see so many cultivars come down with Rust.  One bright spot, however, was that many of my Dip seedlings were resistant even under extreme Rust conditions. Very few Tets showed any Rust resistance, so I decided to work with Dips. My direction from 2002 on, to breed rust resistant seedlings, started with diploids that were already generations away from their named parents.”

Ron working with his daylilies

So to more fully understand the lines that Ron has developed, I felt it important to find out what those diploid seedlings he was working with had originated from. This I felt was important, as I am of the opinion that rust resistance factors are heritable and I had noted in various rust research evaluations and surveys, that certain family lines consistently popped up. So I enquired about the origins of the dip seedlings that Ron began his resistance breeding from. Here is his informative reply.

The most influential daylily in my program was Carpenter's MERLE KENT MEMORIAL, which unfortunately is a tender evergreen.  What I did with that plant is representative of the way my seedlings have been developed.  I selfed MKM and the result was a very wide range of colors, all with perfect full flat form…here are the hybridizers that I patronized in the early days, in order of importance: Jack Carpenter, Brian Millikan, Fran Harding, Dave Talbott, Tom Wilson, Lee Gates, Patrick Stamile, and Pauline Henry.

“The purple line was from CRAYOLA VIOLET (Stamile) crossed with BE MY VALENTINE (Fran Harding).  Some of these seedlings were rust resistant. I still use (some of) those seedlings to maintain rust resistance. (I will have more to say about some of these seedlings later in this article, but for now be sure to note Crayola Violet, as it pollen parent is Super Purple, and the Super Purple family, including many cultivars descended from SP often show up amongst daylilies noted for rust resistance in many surveys and evaluations).

The red line started with DENA MARIE, also a Carpenter, x ROSIE PINKERTON (from Millikan). The yellow line started from Pauline Henry's SILOAM GREEN STRIPE x Tom Wilson's EMERALD SPLENDOR. My white line was primarily a MKM self x some of Brian Millikan's whites.  The oranges are combinations of the red and gold/yellows, and many reds were similar combinations of cool and warm colors.  Mixing my own colors in the burgundy/red/gold/orange range proved to give better results than purchasing a "red" and trying to improve on it. My pinks with red eyes came primarily from Carpenter's PEPPERMINT DELIGHT. THELMA ELAINE AND ANGEL RODGERS from Tom Wilson produced pinks with moderate rust resistance.  Rose-colored blooms came from Fran Harding's WILDERNESS FAT MAMA. I did not purchase any new named cultivars until 2009, when photos of patterned daylilies caught my eye and I invested in some of Carpenter's patterns. I have continued to purchase named cultivars yearly since then, always just for the patterns, and I continue to remove them from the hybridizing program when they show intolerable defects.” 

It is very important to note that Ron points out that not all of the early cultivars used showed rust resistance, nor were all of his diploid seedlings, at the time this work began, highly rust resistant. “My original diploid breeding stock, at the time of the arrival of rust, was not rust resistant (i.e., immune); it was just more resistant than the tets.  I bred the best to the best for generations before I saw plants that could actually stay (completely) rust-free. When I saw that I had a seedling that exceeded the performance of the parent, I removed the parent from the breeding program and continued on with the seedling.” This is important to note, because far too many breeders out there believe that to breed for rust resistance, you must have all cultivars be one-hundred percent rust free, but my experience with breeding for disease resistance suggests that one can start a program with less than 100-percent resistance. In fact, it would be very rare to even be able to begin a program in such a way. Where one should start is with the most resistant cultivars they have or can find and select from there, working to concentrate resistance factors into ever-greater levels of resistance.

Ron working in his seedling bed at peak bloom

At this point, I want to make a distinction between rust resistant and rust immune. A plant that is resistant to rust may show some rust, but will not become completely covered in rust, and the levels of resistance vary from cultivar to cultivar. In my opinion, any cultivar that shows some resistance can be used to breed up higher resistance, so long as it is bred to equally or more resistant cultivars. Immunity, on the other hand, is when a cultivar never gets any rust. While this may be the final goal, not all cultivars that never show rust will have true, genetic immunity, as there is some evidence that some such cultivars may be the result of environmental conditions keeping rust at bay, rather than true, genetic immunity. While it may be advisable to use some seemingly immune cultivars in breeding for rust resistance, especially if they have been consistently immune in various locations or for many years of rust exposure, not all cultivars will truly be immune or pass on immunity to their offspring. So in breeding for rust resistance, the notion that only the immune can be used is inaccurate and Ron’s program points this out.

Now I would like to take a moment to look at Ron’s resistance rating system, to give us a little bit more insight into his process.
I have rated my plants as:
RR (Rust Resistant, or 10) – these never get Rust even under extreme disease pressure.  
HR (High Resistance, or 9) - might get a few pustules on old leaves late in fall.
MR (Moderate Resistance, or 6-7-8) - plants that carried some Rust throughout the season or only in the end of season; this level of Rust didn't hurt overall plant performance or looks. 
PR (Poor Resistance, or 1-6) - plants where Rust started early and took the plant down or damaged it badly.  (I cull Poor Resistance immediately unless they are very unusual.)”
It is important to understand, that in some instances, Ron has used plants from every rating level, but always with a focus toward using one very resistant parent. He feels, and I would tend to agree, that if rust resistance is genetic (as it almost certainly is) and heritable, a plant showing lesser resistance from a resistance-breeding line is possibly carrying resistance factors and can be used should it also offer very unique traits. With that said though, most of his breeding has focused on the RR and HR categories.

So now that we have established some background and the rating levels, let’s look at what Ron has to say about the actual breeding he has pursued in seeking to produce more resistant lines of daylilies. As stated above, once rust first struck, Ron discontinued his efforts at breeding tetraploids and went over to his diploid seedlings to start his breeding efforts over. Those seedlings had been produced prior to the first rust outbreak and showed more resistance than the tetraploids, though very few of them showed high resistance or immunity. Ron was convinced to move forward with those diploid seedlings because, as a group, they showed higher resistance than his tetraploids. Here are some more of Ron’s thoughts on the process of breeding for rust resistance.

We get rust - in the extreme - every year in our garden in Russellville Arkansas. Because of this situation, we changed long ago from hybridizing tetraploids to diploids.  There is absolutely no question that some daylilies have outstanding resistance to rust.  The only true way to control rust is to hybridize new cultivars with the aim of including rust resistance in their genetic makeup.  We have been working to establish rust resistance in our diploid line for over ten years.  We have achieved notable success in producing rust resistant cultivars in a full color range, although some colors are more resistant than others. We started out with a handful, and with rigorous culling, have brought our diploid line up to a 60% RR level.

“I grow from 12,000 - 20,000 seeds per year and I can see the non-resistant seedlings rather fast.  I keep a row of new Southern-bred Tets to get a good strong Rust source, because without a heavy infestation of spores, you really can't make an adequate call on what to cull each season.  I say, bring the Rust on!  Since I needed volume to have any hope of achieving success, I decided to forego tagging crosses as I had in the past, and to make as many crosses and seedlings as fast as possible to create the whole range of colors in Rust-resistant plants. At first I culled all the plants that got Rust in their (seedling) pots. Then I decided to grow all seeds outdoors; it gives a stronger plant and eliminates many would-be rotters.  So now, I collect the seed, stratify it wet in plastic trays, rinse the seed twice a week for 3 weeks, and then plant it out in rows. The seeds generally get planted in late September; two weeks later they are up, which is just in time for fall Rust. In April the seedlings are lined out in blocks 3 x 5 inches apart.

“The previous year's crop is selected and each is given a detailed description on an 8" plastic tag, which is left on until the bloom season is finished.  Then all but the tagged plants are removed, allowing the selections to grow out away from crowding.
By fall they all have new strong growth (as well as) old leaves, which will set the stage for the next evaluation - Rust. Any plants that have (heavy) Rust are culled now, leaving resistant (RR) ones and any that have only light cases of Rust (HR). (I have had to cull some of the most beautiful lilies I have ever seen at this stage.)
The levels of Rust-resistance of the new selections is noted on the flower and plant evaluation tag and only in October are moved into the evaluation block.  Each year I cull, for one reason or another, about 20% of all previous selections in the evaluation block.  Just before the first heavy freeze, all plants in that block are re-evaluated each year and downgraded if necessary.

“The importance of keeping enough Poor Rust (PR) resistant plants to provide adequate infection can't be overstated.

“The year 2008 was the wettest year for us in Arkansas for the past 33 years.  We received 75.45 inches of rain.  It was also the worst year for Rust I have ever seen because of the constant heavy dew.  I had to downgrade some of the 2007 RR (or 10) plants to HR (or 9).  But older plants that I have had for years maintained their resistance code, showing me that there is true resistance, year to year, if properly selected.”

So we can see from this that Ron is following classic selection methods, as any breeder would in selecting for any trait. In selection for resistance, one must expose the individual plants to the problem we are seeking to breed resistance toward, and Ron has done this every year since rust first appeared in his garden by not spraying for rust and allowing it to run through his plants and using it as a culling device. This is a project that anyone who gets rust recurrently in his or her garden can easily pursue, but the desire to do so must be present. Beyond that, I feel there are two more things I wish to point out for this article. The first, which I touched on above, is that you can begin a breeding program for rust resistance by using any cultivars that show above average resistance in your garden. They do not all have to be seemingly immune. The second point is something that Ron states above and that helps me to be convinced that true, heritable resistance to rust really does exist and that he has actually imparted genetic resistance into his lines through selection against susceptibility. Ron states. “But older plants that I have had for years maintained their resistance code, showing me that there is true resistance, year to year, if properly selected.” The importance of this cannot be overstated.

Let us look more closely now at this last point as we close out this article. In any garden that gets rust, there will be some plants that seem immune one year, but are not the next, or in some subsequent years. As noted above, environmental factors can create seeming immunity in some plants. However, such plants, when their immunity is not due to genetic factors, will not produce immune offspring. So how do we determine real immunity or high resistance? It is through replication of results. There are two ways to do this. One is to have many clumps of a given clone growing in many locations, randomly, throughout the infected garden, or in multiple infected gardens. The second is to observe consistent resistance or immunity levels from year to year, over many years, in infected gardens.

So when Ron mentioned seedlings that have been consistently RR in his garden through many years, my interest was peaked and I began to ask many questions about these individuals. While Ron does not have huge numbers of these, there have been a handful of RR and HR seedlings that have consistently scored very high on a yearly basis over many years. I feel that these individuals are very special. Fortunately, Ron recognized their importance, and in many cases is still using them in his breeding as he works toward his eventual goal of large flowered, patterned flowers with very clear colors on vigorous plants with great scapes and branching at about 32” or more in height. While all of these older, proven RR and MR seedlings do not have all of the phenotype points that he desires in his finished line, he has recognized the importance of their long-term rust resistance and so has had the wisdom to keep using them to backcross his subsequent seedlings of more desirable phenotype with. By doing this, he is concentrating their high rust resistance while continuing to select for more refined phenotypes of flowers with improved garden plants. The fact that he has been breeding now for ten years and is only beginning to approach his targeted resistance and phenotype traits for introduction speaks volumes about his commitment to the plants, rather than the desire to quickly produce plants for resale. Such commitment to the plants is admirable.

Ron's seedling bed after end of season culling. This is the same seedling bed in the pictures above!

For anyone interested in Ron’s work, you can see many pictures of his seedlings, complete with some information about each on Ron’s Picasa Web GalleryThere you will find color pictures of many of Ron’s newer seedlings, but a careful review of all the pictures there will reveal some of the older seedlings that have remained RR or HR for many years. Most notable among those are some of the seedlings that go back to 2004 or earlier and have remained RR. Be sure to spend some time finding these seedlings amongst those pictured, as they are very important, both to Ron’s breeding efforts and to demonstrating that long-term resistance is possible.

One of Ron's beautiful lavender seedlings. This is one of my very favorites of his pre-2009 seedlings. To see many of Ron's pre and post-2009 seedlings, be sure to check out his web album, by clicking here or using the provided link in the above paragraph.

In the next installment of this series, we will look at the direction Ron is heading with his breeding, both for resistance and phenotype traits. Ron is a perfectionist, and while I might have introduced some of his RR older bridge plants, he is insistent on achieving his vision of resistance and phenotype combined before he makes any introductions. If only more breeders had this commitment to perfection!

To go to Lessons in Persistence: Part 2 CLICK HERE.