Using Stress as a Selection Tool

Using Stress as a Selection Tool

In breeding any plant or animal, we often have the tendency to pamper our precious offspring, especially if we only have a few or they come from expensive and rare parents. There are instances where this may be warranted, but they are few and far between. If we are talking about a very rare parent or narrow gene pool, or if we only have a few offspring, we may pamper those few in order to get them to reproductive age and increase the rare stock. In time though, as numbers increase, our goals should shift toward selecting for the most vigorous and hardy individuals within the gene pool to continue increasing the numbers of that particular family line. In the vast majority of instances though, pampering does not help the family line nor is it helpful in gene pools that have large numbers of individuals. In fact, in those instances, it can be detrimental.

In breeding, we usually understand that the things we actively select become concentrated into our gene pools, but we often fail to realize that those things we ignore and do not actively select can become imbedded in our gene pools just as easily. I refer to this as passive selection, in order to contrast it to active selection. In active selection, we actively select for certain traits, generally seeking to increase them. With passive selection, we are not actively selecting for a trait or traits, but we accidentally select those traits through ignoring them or not creating conditions that reveal them.

An excellent example of this is crown rot. We know there is some genetic aspect involved with this deleterious trait, as crown rot is prominent in particular family lines. So when we use those family lines, we run the risk of creating more cultivars with the propensity for crown rot. In order to select against the crown rot trait, we would want to expose seedlings to the conditions that are conducive to crown rot, in order to select those seedlings for further evaluation that do not show the trait and then to also eliminate those seedlings that do show crown rot. 

I have found two techniques to test against crown rot. One is to grow seedlings in fairly moist conditions and the other is to divide and replant seedlings in hot, moist conditions. By doing these things you can identify those that have a high propensity to crown rot and eliminate them (if they don't actually eliminate themselves for you). 

This picture shows some of my seedling growing tubs. Note that the drainage holes are drilled 1/3rd of the way up on the sides, rather than on the bottom of the containers. This is done for two reasons, both involving the creation of a water reservoir. The first reason is that this reservoir ensures there is water retained there so that in very dry weather, supplemental watering is not necessary on a constant basis. The second is that in wet weather, and through much of the winter, the seedlings are kept constantly moist. In hot, wet weather this will activate rot in those families that are susceptible, while families that are not susceptible show no crown rot. This added water stress can eliminate rotters, but in those individuals that have no susceptibility, it actually increases the rate of growth, thus serving two purposes.

However, I frequently hear people talk about how they only move their seedlings late in the year when it is cool to "avoid rot". This is a way to avoid seeing rot, but by not seeing rot, you can not make selection against the trait. You are then passively selecting toward the propensity to rot because you are not actively selecting against it. There is always the chance that the seedlings you select for further breeding or introduction won't carry the rot trait, but there is just as high a chance that you will select the worst rotters and if you don't provide the conditions to reveal those traits, how will you know?

Rust is another excellent example. There are genes for resistance to rust in daylilies, but a great many breeders spray to suppress rust from going into the spore phase throughout the year so there is no way they can be selecting for those individual seedlings that show higher levels of resistance or immunity to rust.

In both instances, the breeders undoubtedly feel they are doing the right thing and taking the best possible care of their plants, but the "best possible care" of the individual plants is not always the best possible situation to select the best possible plants in the next generation, though it might allow you to select "the best" flowers, if that is your only focus. In short, the best interest of the entire gene pool is not always the same as the best interest of the individual plant.

In breeding it is very important to remember this. If we pamper our plants those individuals may flourish, but we may thus be compromising our gene pool by not having proper situations by which to select for important traits such as plant vigor or disease resistance. The only way we can really select for such traits is by creating (or allowing) the conditions that reveal the plants with the best concentration of genes for the desired traits.

So let's take a minute to think about some of these (rather obvious) points.

1. If you are concerned about drought tolerance, don't water your seedlings (and maybe your breeders too).
2. If you are concerned about disease resistance, expose your seedlings and breeders to the diseases that concern you and the conditions that accelerate the disease(s).
3. If you are concerned about flowers opening on cold mornings, grow your seedlings on the Northern side of your house or garden, or in a cold-pocket microclimate.
4. If you are concerned about water spotting, then overhead water when scapes are flowering.

While this area of my hybridizing garden may look lush, it is in fact very dry. A south facing slope, the soil is a mixture of sand and orange clay with a very thin layer of topsoil. I use this particular area to test out potential breeders and seedlings for drought tolerance. This area is too far away from the hose for convenient watering and I am highly unlikely to actually carry buckets of water very far. In the drought of 2012, I was able to produce over 7,000 seeds from this area because the cultivars I focused on there have proven themselves fertile in full sun and dry conditions over several years. Not surprisingly, the seedlings from those cultivars tend to also set seeds well in full sun and dry conditions...

These are just a few ideas. There are many, many more examples. Think of the traits (faults) that you want to work against and then think about the conditions that seem to bring out or accelerate these negatives and how you might expose your plants to these conditions in order to reveal those with the weakness, as well as, and most importantly, those that do not display the weakness. Select the later to move forward with!

I was recently watching a Youtube video where Dan Heims of Terra Nova Nurseries speaks about how new perennials enter the market. It is fascinating and I recommend you watch it, but I was particularly excited by one comment he made in regards to breeding new plants. He states that he starts all breeding projects by asking the question, "What is wrong with this plant". He says that he is looking at the problems in order to solve them through selection. His story about breeding pulmonaria for mildew resistance is fascinating and every daylily breeder should take head to the successful techniques of this master breeder. Below is the link to the Youtube video of his talk. Enjoy! 

Dan Heims, Perennials from Around the World

Thoughts and Observations: Inbreeding

Thoughts and Observations:

Inbreeding

Brian Reeder

*I wanted to note at the beginning of this post that I am writing one post this month for the blog, which will also be used for my monthly article in the Bluegrass Hemerocallis Society newsletter as well as for Exhibition Poultry E-zine. This article is a general article on breeding techniques with a focus on inbreeding and is fairly equally applicable to animal breeding and plant breeding. I may use specific examples of a particular plant or animal, so if you want it to specifically apply to your plant or animal, just change the name in your head to what you are working with. The underlying principals are the same, with only one exception - you can't self-mate (most) animals as you can with many plants.

Breeding in many ways is as much an art as a science. Science (genetics especially) is always a valuable tool in the breeder’s toolbox, but I always like to point out that our ancestors created the vast majority of domestic plants and animals long before there was any knowledge of genes, beyond perhaps the most cursory understanding that "blood will tell". The art of breeding, like any art, has a lot to do with individual taste. What I may adore, you may despise and vice-versa, but in the end, we all use basically the same set of techniques to produce our desired outcomes. 

There are really only so many approaches to breeding that one can take - outcrossing, backcrossing, line breeding, inbreeding, etc. We all use these techniques to one extent or another. It is interesting to me that in some circles though, one or the other of these techniques have taken on a superstitious veneer of 'taboo' status. For instance, in ornamental plant breeding, we see a majority of outcrossing and little inbreeding (or at least little that is openly admitted), generally due to the old notion that inbreeding is "bad". In animals, we see little outcrossing (again that is openly admitted) but lots of inbreeding, due to the equally superstitious notions of "crossbred" and "mongrel" or "un-pure" as opposed to "purebred". Each set of suppositions limits the breeders in those respective arenas.

There are no good or bad breeding techniques. There are only breeding techniques, and in some instances, each is of great value while in other instances, any given technique can be disastrous. It really all depends on the individual situation and the desired outcome. Inbreeding particularly gets a really bad rap, and there are some very negative potentialities to inbreeding if it is not applied carefully.

Inbreeding is used to concentrate genetic traits. It is that simple, and inbreeding will concentrate whatever is there, good or bad. Inbreeding can be especially valuable in determining what recessive genes are lurking in a particular population. If there are bad genes, deleterious genes, hidden in a population you can bet that inbreeding will bring them to the surface, but there is also no better way to determine what bad genes are lurking in a particular population. On the other hand, there is no better technique than inbreeding to determine what good genes are lurking in a population. As well, inbreeding can reveal how extreme a given trait can become and if a trait that is neutral in lesser expression will become dysfunctional or deleterious in higher concentrations. 

I have used inbreeding a great deal in strain development both to concentrate desired traits and to reveal undesirable traits in both plants and animals. I always approach inbreeding with a very careful eye, quick to note when there are undesired traits and I am equally quick to eliminate a line or discontinue inbreeding when it becomes clear that there are many deleterious factors carried in a line. However, in some instances, inbreeding for generation may reveal no deleterious factors and in such instance, may only concentrate and strengthen the positive traits of the lineage.

In my work with daylilies, one of the first tests I like to run on a given cultivar is to self it. That is, breed it to itself. The first thing I am looking for is if the cultivar is self-fertile. The next thing I am looking for is how clone-like the seedlings of selfing may be, or how much variation they show. The former indicates a great deal of homozygosity and/or many dominant genes, while the later indicates heterozygosity of some or many genes. In the case of the former, I will then know that I may not be able to create anything very different from the parent in an F1 outcross, and so it becomes a producer of bridge plants for further breeding. If the plant is highly heterozygous for many traits, then one can expect to produce a wide range of phenotypes in the F1, and feasibly something different enough from the parent or unique enough to potentially be an introduction in its own right. However, I still consider such a cross more for the production of breeding material than for introductions, but that is my focus anyhow. For me, introductions are really the last thing on my mind with any cross at this time, while the production of new and improved breeding materials is always paramount in my efforts.

Chickens can't be selfed, but they can be crossed with full or half siblings and they can be backcrossed to either parent. To me, this is the only really tight inbreeding that is likely to reveal the hidden traits I wish to evaluate. The breeding of cousins, for instance, while inbreeding in the broadest sense, is to my way of thinking more in line with line breeding, and less pure inbreeding. While the purest inbreeding in plants is selfing, sib x sib matings as well as backcrossing to either parent (or aunt/uncle, grandparents) are useful techniques of inbreeding and can be equally as revealing for strengths or weaknesses as selfing. Backcrossing is particularly suited to revealing the hidden details of one or the other of the parent lines. The biggest pitfall with inbreeding comes when people get so tunnel-visioned on a particular trait that they ignore problem genes, sacrificing viability for phenotype extremes.

To put it more bluntly and in more common language, if you have bad traits in a line and you inbreed, you are likely to destroy that line as the bad traits become more and more concentrated. Conversely, if you have good traits, but no bad traits, inbreeding can concentrate those good traits and make them even more prominent. In reality, most lines have some of each, and so inbreeding must be approached cautiously and with care, being always aware to stop inbreeding when bad traits become glaringly obvious, no matter how "good" the line looks otherwise.

In animals, any given breed or line is likely to be the product of some inbreeding already. With animals, inbreeding can very quickly cause a lot of problems, and only the most vigorous and hardy lines are usually good subjects for intensive inbreeding. However, inbreeding is much more common in animals, thus the many problems we see in many, many breeds of domestic animals. The only instances where I would encourage intensive inbreeding in animals is where there are no obvious deleterious factors present (so, so unbelievable rare!) or where you have outcrossed and are working to make a new line, variety or breed and wish to concentrate good traits while working very hard to eliminate bad traits.

Plants may be another thing altogether, though. In daylilies, for instance, there are no know "pure" lines, as we see in so many domestic animals. Almost all daylilies will be heterozygous for a lot of alleles and are the results of much outcrossing, starting with the hybridizing of species about a century ago. It is my opinion that daylilies may be good candidates for some careful inbreeding, in some cases. Now we don't want to get carried away (as a cautionary tale, just look at what the animal people have done to their charges through "pure-breeding"!!), but there is much that careful inbreeding may accomplish. Of course, we want to be ever vigilant about inbreeding depression and the concentration of bad traits, but since most daylilies are so heterozygous, I doubt we are on the precipice of inbreeding collapse, as are so many domestic animal breeds. I would note however that certain traits seem to be questionable in daylilies already, so concentrating those traits may be unwise. For example, think edges so profuse they don't allow the flowers to open well or weak scapes with oversized flowers, etc. Even without intensive inbreeding in daylilies, there are problem areas and these need to be admitted and faced so that they don't get concentrated into 'ruined finery', so to speak.

To consider inbreeding in daylilies, let us consider an example. Let us say that there is a new, novel trait that has appeared in daylilies and we want to perpetuate that trait and perhaps even intensify it. If the trait is strongly dominant, then we can just cross to most anything, selecting those that express the trait most strongly and those, which combine the trait with other traits for new, novel phenotypes. But if the trait is recessive, then outcrossing it willy-nilly, crossing "pretty to pretty", is liable to give us very little that we want in the F1. Granted, that F1 can be interbred to bring the trait back out, but I have encountered very few daylily breeders who are going through generations of bridge plants to get to the desired outcome. Most seem to chuck the whole batch if they don't have an intro in the F1, though that doesn't apply to everyone and there are already some notable examples of just this type of inbreeding in the history of daylilies. 

So viable options for breeding such a trait would include selfing, backcrossing to either/both parents and/or creating an F1 through outcrossing and then interbreeding those F1 or backcrossing the F1 to the parent with the desired trait and/or the grandparents that produced the trait in the parent showing the trait. While I would hesitate to do anything other than outcross and interbreed the F1 if there are problems in the parent with the new trait, I would also want to look at its parents to see how strong each of them are. 

Since daylilies are highly heterozygous (generally), a weak plant or one with a given problem may not be concentrated for that trait. If the parents of that plant are easily obtainable and/or much is known about them, you may find that the problem trait is a dominant and may be heterozygous. If that is the case, then selfing or any other inbreeding may allow you to select out the desired trait in combination with other good traits and to eliminate the undesired bad trait or weakness. Genetically, things are not always as they seem, or more technically, phenotype is not always a full indicator of genotype. It is only through some experimentation that you can really know what is what genetically, the true breeding value, and thus have a fairly clear idea of the real limitations and strengths of a given cultivar or seedling, and one outcross that produces 10 or 15 seedlings really isn't going to tell you much.

The instance that I describe above is very analogous to animal breeding where outcrossing has been done (except for the selfing, of course) and a new combination of traits is being sought; i.e., a new breed or a new variety of an existing breed. Pure breeding is rather unique to the animal people. Can you imagine the ornamental plant breeder that would be striving to produce a cultivar that never threw offspring that looked any different from itself? (While this does occur in some commercial plant breeding, especially with some of the old, true breeding food plants, it is by far the exception in breeding for ornamental plants.) The thought is laughable and ludicrous, generally speaking, but it is a deadly serious affair to animal breeders where the brainwashing tends toward the notion that each individual of a given breed or line should look like identical twins or clones. Domestic plant breeding is much more in line with the notion of 'landraces' in animals. I think there is a lot to be said for landraces, especially the fact that it is nearly unheard of to find a landrace that is so concentrated for deleterious traits that their very existence is questionable. With that said though, there is a place for inbreeding in both the plant and animal worlds but the practice must be approached with caution and an understanding of the potential pitfalls, along with an eye for the slightest indications of problems. When used wisely, inbreeding is an indispensable tool for the breeder.

What Makes A Good Daylily?

What Makes A Good Daylily?

“What makes a good daylily?” This is a question I often ask myself, and I have been thinking on it for many years. I also often ask other daylily people this question. I get some interesting answers, but in most cases, the answers people give usually starts with the flower and only concerns the flower. My personal answer starts well below the flower with the roots.

The root of a plant is the base from which all the rest of the plant stems. The ability of the roots to uptake nutrients is the source of much of the ability any given plant has to perform. So why do we rarely hear much discussion of the roots of daylilies? When digging and dividing, or planting new divisions, we all encounter the roots of our daylilies. It is true that they are hidden from view on our plants in the garden or field, but we still see them with some frequency, so why do we not think about them much? Of course, I suppose we could often ask this same thing about every part of the daylily except the flower. It is a rare daylily breeder indeed who is giving a lot of focus to any part of the daylily other than the flower.

Since we have all seen the roots on our daylilies when we plant them or divide them, do you pay much attention to the variations in daylily roots? This is something that I pay attention to. I have noted a wide range in root structure from small rounded roots to long, thin roots to long, thick roots to roots that produce rhizomatous growth. Some daylilies produce new fans that are just barely connected to each other and are easily divided, while some grow all their new fans from a solid main crown. The roots of each crown type can be very variable.

From a grower’s perspective, is there much difference in performance based on the roots? Well, in my experience, the answer is yes. I have noticed that the cultivars with small, short roots often are small plants that may not have very vigorous performance (except in some very small dwarf forms). Plants with long roots with lots of feeder roots coming off the main larger roots seem to show a very vigorous performance. Rhizomatous growth may not be desirable for everyone, but for covering an area, it is very desirable. However, in a garden situation where contained clumps are desired, rhizomatous growth is very undesirable. I would note though that I have never seen a rhizomatous type that was a weak plant with poor performance.

The variability of roots and crown structure is very interesting and may be something to which more attention should be paid. It is my experience, for example, that dividing a clump where each fan is separate from the rest in the clump is much easier than dividing a clump where every fan grows from one main crown. While this may not make much difference if you are only concerned with growing the plant in your garden, for breeders and sellers, this can make considerable difference. I would note that I have personally noticed much more crown rot in those types that have one crown from which all the fans grow and that must be cut apart to make divisions, than in other types. The cutting apart of the crown must be much more stressful when dividing than simply pulling apart individual fans.

So should a breeder never use a plant with a joined crown or that shows small roots? I would say no, because sometimes there is a desirable trait that is found with undesirable traits, and the art of the breeder is to recombine desirable traits while eliminating undesirable traits. I have a few breeder plants that show various undesirable traits. There is no way around this, especially when a given plant may have a very unique trait or offer some other very desirable traits. The key to using such plants is to mate them with plants that show good balancing traits. So a plant with one solid crown could be mated to a plant that multiples through making multiple individual fans and the offspring could be deselected based on solid single crowns. In another example, a plant with smallish roots that is not a very vigorous grower could be mated to plants with strong roots and vigorous growth and the offspring deselected for small roots and weak growth. Also bear in mind that not all goals are achieved in one generation, and some traits can be recessive in nature.

One example I would site is Grand Masterpiece, which is a lovely dark purple diploid with very strong rust resistance, but the roots are small and the plant is not a vigorous grower. However, Grand Masterpiece (either the dip or the tet conversion) can produce offspring that are much more vigorous with much stronger root growth. As well, Super Purple, one of the parents and of Grand Masterpiece, is a vigorous plant with strong root growth, so the evidence in that case suggests that Grand Masterpiece has the genes to produce more vigorous offspring and better root systems, as evidenced by some of its offspring. Would I recommend Grand Masterpiece as a garden plant? No. Would I recommend Grand Masterpiece as a breeding plant? Yes.

It is important to understand that some of the things we may grow as breeders are not desirable garden plants, but carry special genes that make them useful breeding plants. It might be a good idea to be very clear as to which cultivars are which, as average gardeners who accidentally buy a highly-touted breeding plant expecting it to be something special may think twice before they invest in more highly-touted daylilies. The public will often equate high price with special performance or garden value, but it is far too often the case that the most expensive daylilies aren’t great garden plants. Instead, such plants may only be of use to a breeder who knows how to select proper mates for them. We need to be careful not to turn off potential gardening customers. With rust on the horizon and more people having less disposable income, I am afraid that we cannot afford to turn off potential gardening customers. I am all for breeder’s plants, but they should be noted as such. I have a few very expensive plants that have many undesirable traits and I would never recommend them for the garden, but I greatly value one or two traits that I am extracting from them, struggling to recombine onto much more desirable garden plants.

When looking at what makes a good daylily, I am first and foremost concerned with how any given daylily performs in a real garden, an average garden. Most average gardeners don’t have the resources to give any particular plant much pampering. It is true that many expensive modern daylilies perform beautifully when kept under shade cloth with automatic irrigation and feeding of very exacting fertilize formulas while growing in expensive and refined growing mixes, but those same daylilies often look terrible and perform even worse in a real garden situation with blazing sun, little or no irrigation or fertilize and average (or worse) soil. So when I look for a good daylily, I am first and foremost looking for a daylily that thrives in average, real-world garden situations.  Roots may play a very big role in that, but genetics and selection will play a big role also.

We can’t expect plants bred from weak stock to come out strong and vigorous in the first generation, unless, as in the example above of Grand Masterpiece, we know something of the parents of the cultivar and then mate the cultivar to vigorous plants that balance its weaknesses. Even then, we may have to go to a second or third generation to combine all the desirable traits of performance and flower that we wish to see. We also can’t expect plants bred in highly pampered conditions to show real-world hardiness, though some of them may. For a breeder who is interested in producing plants that show great garden traits that will satisfy the needs of real-world gardeners, we would probably be best served to grow our seedlings in conditions that expose the weak and highlight the hardy and vigorous.

One thing that breeders should always keep in mind is that selection follows exposure. A negative trait may only be exposed when conditions are present to allow the negative trait to be seen. However, some plants are weak or show major flaws in even the most optimum conditions. Can you imagine how poorly these plants do in average conditions? While such a plant may be introduced for breeding purposes, I would always hope that breeders are very clear about its negative traits so people who just want a nice garden display will avoid it. You may say, “Well, surely the price will keep such people from buying it”, but I fear that is not always the case, as too many people equate expensive with high quality.

So long as we pamper our plants and don’t expose them to some bit of adversity, it can be very hard to know what has good garden potential and what is only good in optimum conditions or for breeders. While many breeders may want to give their cherished specimens the best possible care (and who can fault them? I certainly don’t!), there are many clever ways to test any seedling without neglecting the entire garden. One is to simply remove one fan from your select seedlings and to grow that fan in less than idea conditions to see how it performs. Another is to send divisions to friends who love to garden but do not have great garden conditions or refuse to pamper, and see how the plant performs in their conditions. This may slow down the rate of introduction of new seedlings, but that might prove to be a very good thing. I know more than one breeder who has told me that this or that got introduced too quickly and proved to be a poor plant on down the line after introduction. In a few instances, I know of cultivars that only got introduced after several (or even many) years because of how the plant performed over time.

“Good” daylilies will vary based upon the definition of “good” that each person uses. It seems that for many, many people “good” only concerns the flower. For me, “good” is first and foremost about the plant, its ability to thrive and multiple, and its ability to do so without being pampered. While this may seem simplistic to most daylily fanatics, this is also the greatest concern of most average home gardeners. If the daylily is to remain a popular garden plant and the world at large is to learn of the wonders of modern daylilies, we need to produce daylilies that will perform well when average gardeners grow them and we must to begin to respond to serious disease issues such as rust through testing, breeding and selection. Otherwise, home gardeners are likely to stick to a few varieties they know to be reliable (Stella, Happy Returns, Little Business, etc) and all of our marvelous advancements will go unnoticed, perhaps in time even lost, with huge potential markets untapped.

Pod Parent Selection

Pod Parent Testing to Determine Breeding Qualities of Individual Clones

Brian Reeder

     As a beginner to daylily breeding, I am in no position to speak of details concerning daylily phenotype segregation and combinations, but as a long-term and experienced breeder, I am in a position to speak about breeding and strain formation. I want to state for the record that what I am going to outline below is simply what is important to me. As a breeder, it is up to you to determine what matters to you. The program I am presenting below is simply a stage in my over-all breeding goals. This phase is the foundation of strain development. While I would, as an experienced breeder, recommend strong attention be given to this phase of strain development, I am not saying you should do this. How you approach your breeding is up to you. I hope what I present here gives you good food for thought about strain formation.

In the beginning stages of strain formation, I am most interested in laying down the most basic traits that are required for the best health, vigor and reproductive ability of the given organism. These traits are paramount, as they lay the foundation upon which phenotype traits can then express and reproduce. That is not to say that the physical traits of an organism necessarily cause specific phenotypes, but rather that the heritable health, vigor and vitality of the organism, in part, determines what is possible in the way of phenotype expression. In my own way of looking at a given individual or strain of an organism, I see the advanced phenotype traits as ‘window-dressing’. In other words, they are simply the decoration we apply at the end of a long building process.

To draw this metaphor out a bit, I can say that I wouldn’t want to hang drapes until I had a house built. I would have laid the foundations, framed in the walls and built a roof, then finished out the inside of the structure before I did things like hang drapes or put up pictures, lay down rugs, paint or put up wall paper. Over my three decades of breeding many different organisms, I have come to see phenotype traits as such finishing touches. Much like house décor, these things can be changed. If we are looking at houses to buy, choosing the one with the wall paint color you like the best but a severe termite infestation would not be wise. You would look for a sound house and then paint the walls any color you want. I approach strain formation in the same way, looking for basic, foundational traits upon which to attempt to build a sound strain for later phenotype selection.

To apply that to daylilies we might say that a weak, hard to grow plant may not give the optimum flower performance, that which we might consider optimum for daylilies of similar ploidy in general. Or conversely, we could say that a strong plant may be able to give more energy to flowering. However, I have seen instances of fast growing daylilies that do not produce many scapes, so it is clearly more than vigor that creates optimum scape densities and bud counts or reblooming traits. It would seem such traits have their own genetic basis and thus we may see a slow growing plant have a high scape density or bud count, which is desirable, but simply is not be a strong plant. However, in some instances, we see a combination of vigor with scape density, high bud counts and/or rebloom. In my experience with collecting daylilies for the last thirty + years (I am 43 at the time of writing and have been growing daylilies since I was a young child) I have noticed instances of stand-out cultivars, in both ploidy levels, that show strong, vigorous growth and give excellent performance for scape density, bud count and/or rebloom. In short, such individuals are outstanding plants that are a joy to grow and reward you for the least bit of effort.

A further consideration is that the expression of such desirable traits can vary from cultivar to cultivar to some extent depending upon environmental conditions. Even environmental variations within one garden can cause variable expression of phenotypes in daylilies. So what I consider my best examples for the combinations of all these traits may not be so in another environment. With that said though, there do seem to be some cultivars that are reportedly vigorous and show good phenotype expression in a wide range of environments and even seem to be able to pass such traits on to their descendants.

At this point in my breeding program, I am at the beginning of strain formation. I am not choosing one particular ploidy, form, or color of daylily to focus on. My only focus is finding strong plants with good plant traits that are strongly fertile and are able to pass their good plant traits to their descendants. My first focus is to locate and identify my “females”; i.e., the most reliable pod parents for producing both good quality seed and good quality seedlings showing the same strong plant traits as their pod parent. There are many phenotype traits that I admire; forms, colors, etc., so I am making an effort to bring in a wide range of phenotype traits. I am working with over three hundred cultivars, about 2/3 diploid and 1/3 tetraploid, as well as a few species and species clones. I have cultivars that I have grown for nearly thirty years, to newer cultivars that have only been added in the last year or two. I have spent considerable energy in the last few years researching the traits of various daylilies that interest me, both for phenotype and plant characteristics. I have recently added some cultivars due to their advanced phenotype traits for form and/or color, while others I have added because of numerous reports of excellent plant traits.

So to begin the process of forming strains, I spent a great deal of time thinking on what traits mattered most to me. The list of traits below is arranged in the order to which I assign importance to the given trait. Your order of importance may vary from mine. While you are welcome to use my list if you wanted, I certainly don’t expect anyone too. These are only the traits I consider to be of importance and the order is only the order that I place value on the traits. I would say though that the basis of my trait-ordering is based on three decades of intensive breeding and the recognition of the commonality of certain of these traits across all the organisms I have worked with, as the basis of a superior strain, in both plants and animals.

Here is the list of the traits that I am looking at with each cultivar’s seeds/seedlings. My first major breeding season was 2010 followed by 2011. In 2010, I purchased many new cultivars and purchased seeds of many interesting phenotype crosses. These were to be used to select a small number of vigorous an/or unusual phenotype combinations for use as bridge plants to bring in given traits to the formation of strains. I also produced seeds on a handful of my oldest and most vigorous cultivars. In 2011, I pollinated every clump I had, old and new, in order to test all of those many cultivars against the list below. Each cultivar was generally bred to several different pollen donors but the pedigree, for the most part, was only of the pod parent, as this round of breeding was to find the best seed producers, with the best germination and seedling growth. Only later will consideration of phenotype traits be given, and those identified to have the best expression of the first few traits on the list will be taken on into pedigreed breedings in later seasons, as part of more orchestrated strain formation.

1. Ability to set seeds

2. Quality/quantity of seeds

3. Germination rate of seeds

4. Growth of seedlings

5. Percentages of foliage type from given cultivar (i.e., can cultivar produce dorms)

6. Health and vigor of foliage of seedlings

7. Average time of first bloom of seedling group from given cultivar

8. Quality of scapes of seedlings from given cultivar

9. Quality of flower (sun resistance, lack of spotting, clarity of color, etc.)

10. Potential recessive genes carried by cultivar

11. Range of phenotypes in seedlings of given cultivar (flowers)

12. Possibility of Rebloom

With the full list given, I would like to look at each one briefly below.

1. Ability to set seeds

This one is obvious. If a given clone does not produce seeds, it cannot be a pod parent. Some of these plants may be useful pollen parents. Cultivars that are totally infertile one season may show fertility in later seasons. However, my focus is to find those cultivars that show strong seed production regularly.

2. Quality/quantity of seeds

All seeds are not created equal. Some plants produce copious seeds, but those seeds germinate poorly. Other cultivars produce few seeds, but they show good germination traits. It would then seem that these are separate traits that can recombine. The recombination that I most desire is both good numbers of seeds and seeds that store and germinate well. In other words, seeds which are forgiving and germinate in spite of my care. A good example of this in tetraploids is Custard Candy, in my garden.

3. Germination rate of seeds

Once germinated, survival rates of seedlings seem to vary from cultivar to cultivar. I am paying particular attention to those cultivars that germinate well and then also show high survivability rates of the young germinated seedlings. Again, Custard Candy shows this trait in my experience.

4. Growth of seedlings

Once germinated, even when germination rates are equally high, some cultivars show better growth than others. Those that show the fastest growth are noted.

5. Percentages of foliage type from given cultivar (i.e., can cultivar produce dorms)

I prefer dormant foliage, so when I am using an evergreen or semi-evergreen, I am interested in the cultivar’s ability to produce any dormant offspring. My goal is to work toward only dormant foliage, with hard dormant being the most desired outcome. In some instances, I will probably have to use semi-evergreen or evergreen cultivars, so known dormant carriers could potentially be used to produce dormant foliage in their offspring. I prefer to cull seedlings for foliage type the first winter into spring before I have seen the first flowers.

6. Health and vigor of foliage of seedlings

Nice, bright green to blue green to reddish foliage all look nice to me. Yellowing, chlorotic, streaked, or dying foliage is not attractive to me. Those cultivars that show consistently the nicest foliage, as well as some level of frost resistance, will get special focus. Those that have scored well in all the previous points and show desired foliage traits will gain special focus in my strain formation. To me, this is an extremely important point.

7. Average time of first bloom of seedling group from given cultivar

It has been suggested to me that in breeding for the reblooming trait, those individual clones that bloom earliest, from stock known to express rebloom, show a higher likelihood to show reblooming traits themselves.

8. Quality of scapes of seedlings from given cultivar

I don’t like floppy, weak scapes. I do like strong, thick scapes that can easily support their flowers, more than one at a time. Those plants that show good scapes and also produce seedlings showing good scapes will take a special place in my foundational strain building.

9. Quality of flower (sun resistance, lack of spotting, clarity of color, etc.)

Here finally we reach flower phenotype traits. If all other criterion before this shows good scores, then the flower is considered. Like most everyone, I like sun and rain-fastness. I like a range of forms, color, and patterns. Distinct colors are nice, and clear, clarified colors are even better, but some of the smoky or grayed cultivars as well as brown cultivars, are very attractive to me. The substance of the flower is very important to in my opinion, as is the ability of the flower to open wide. I don’t especially care for more trumpet shaped flowers, though I do like some of the pleated or cristate-formed cultivars. Cultivars that have scored well on all previous criteria and show interesting and desired flower traits will then be given special focus in base-strain formation.

10. Potential recessive genes carried by cultivar

There are certain genes that I value that are thought to be recessive (classical Mendelian recessives in diploids). The ‘melon’ factors/clarification factors, reblooming genes, dormant foliage and rhizomatous root growth are all said to be one or more recessive genes. Instances of a parent that does not display the given trait but can produce it in its offspring are to be noted and made use of. Such plants may only be bridge plants, but they can produce the recessive traits that I have chosen to pay attention to. This one is not really so much a selection criterion, except in a few rare instances, as it is a nice thing to know in addition to the previous criteria. Those plants that have passed all the previous criteria well may get used enough to make determination of heterozygous traits.

11. Range of phenotypes in seedlings of given cultivar

This is the ability to recombine traits, i.e., the lack of homozygous dominant traits - this is broadly the general combining ability. Do the seedlings vary widely or are they very much like clones of the pod parent? The later may indicate a high level of homozygous dominant traits. The f1 from such a cross, while looking like the pod parent, may well carry recessive traits of value. Some cultivars produce consistently nice offspring, while others produce a few nice ones and a bunch of not so nice ones. Those that can consistently produce the most surviving strong seedling plants, with other desired phenotype traits, will be given special focus. At this point, a cultivar or seedling that has reached this level, has scored high in all previous criteria and also produces consistently strong seedlings with acceptable flower phenotype traits will move to the highest levels in strain formation becoming major pod-line founders for use in pod and pollen pedigreed breedings.

12. Possibility of Rebloom

This is my final criterion, as it is the last in the cycle of all the traits that will tend to express in the phenotype, in chronological order. Consistent rebloom is something I very much like and is a very desirable trait. Any plant that consistently reblooms and has shown good traits in all the previous categories will be foundational to reblooming strains, which I would eventually like to find in all my strains of daylilies. However, a fine plant, scoring high in all previous criteria, will still be made use of if it does not rebloom. Perhaps in time I would have enough seedlings that rebloom to go over to nothing but reblooming lines, but that day is probably a good bit away at this point.

I expect only a tiny fraction of the cultivars I have to meet many of these criteria, but it is important to isolate those few individuals that come close. These then can be used to build strains that show many desired traits all combined into single plants, both through interbreeding such plants and by outcrossing them to plants with desired phenotype traits in combination with undesirable plant traits. Such an undertaking is a slow process, but I have found that it does allow for the formation of unique strains showing the combination of many desirable traits, in both plants and animals.

Extremes

The Danger of Extremes in Phenotypes

I just wanted to write a few words about the race toward extremes in phenotype that we so often see in the breeding of domestic plants and animals. While this is very common, and might in fact even be considered the essence of phenotype breeding, it can be dangerous to the health, well-being and continued popularity of any given domestic population. This, of course, depends on what the 'extremes' in question may be.

In the case of selection toward extreme examples of hardiness, disease resistance, fertility or adaptability to varied climate/environmental conditions, we would not be seeing something dangerous to the genus, usually. However, when the selection for extremes in daylilies specifically involves larger and larger flowers without the plant underneath them to support them, wider edges with bigger and bigger bubbles/teeth/ruffles that open poorly, pretty faces regardless of the plant qualities, tall/weak scapes that can't support more than one giant flower open at a time (if even the one), extreme flowers that cannot perform in average garden conditions and require ideal conditions or greenhouses to fully express there phenotypes, pretty faces that are not fertile, and plants that are weak in general, that cannot tolerate real-world conditions or have no resistance to diverse conditions including stress factors such as drought, heat, cold or disease, then the extremes can be creating a two-fold danger.

The first of these dangers is that we are creating a weaker plant that cannot thrive (or perhaps even survive) in a real-world garden, thus loosing the public interest. The second danger is the erosion of the genome of many lines that could potentially bottleneck into a dead-end due to health and/or fertility issues, leaving us with only vegetative reproduction (where such plants are able to increase enough to even be multiplied) without any hope of continued sexual reproduction and thus, no hope of new cultivars from those lines.

While what I am proposing here may seem extreme (and we are admittedly not yet at the precipice), and many may see this as only fear-mongering, all we need to do is look at the history of some of the other domestic lines of animals and plants that have been selected for extremes for a long period of time to see such negative potentials made manifest. There are many examples in the animal world and plant world of domestics that have been selected out of existence or to the brink of existence through a focus on extremes at the cost of survivability/reproducibility.

While I am not suggesting that everyone should start breeding for extremes of hardiness or resistance, I would suggest that keeping hardiness and plant qualities in the back of your mind, at all times, is a good exercise for all plant breeders. It certainly can't hurt, and maybe it will help. Maybe, the next time you have a wonderfully pretty face on a poor plant, instead of introducing it, you could just save it for use as a breeder and work to bring its lovely traits onto a better plant? I know many good breeders already do this. Good breeding requires patience, and all good things come to those who wait. While it has been suggested that patience is the reward of patience, in breeding plants or animals, the reward of patience is often a much better end product.

Traits of Daylilies

Traits of Daylilies:

A Consideration of Extreme Points on a Spectrum

As I discuss various trait sets below, it is important to remember that most of these traits will fall along a continuum so that only a few cultivars may fall into the most extreme ends of any given trait set. Further, please bear in mind that these descriptions are only my thoughts and opinions, based on my own experiences in my own gardens. Now, of course, I think my thoughts and opinions are right on target... for me, but they may not be on target for you. A trait that really irritates me may be one that you really like. To each their own! You have to decide what you like or don't like. Clearly, I have made some decisions about what I like and dislike. Please, never take any of my posts to mean that I am telling you that my thoughts are written-in-stone facts, or that I think you should follow my methods or agree with my thoughts. Think of all my posts as my process of thinking out loud, a process through which I am working to iron out my focus for my own breeding program. Please take anything that you like or agree with or want to integrate into your own program and ignore that which does not apply to you or your program. 


It is my belief that there are no "bad" daylily breeders. I suspect that every introduction from a given breeder was beautiful in their own rearing system. There are certain hybridizers that I have many introductions from, yet in no instance has every cultivar I have from any one of those hybridizers been a winner for me (though some have more winners in my gardens than others...). I think that would be a given for any and all hybridizers efforts. They are simply not all going to work for everyone, in every situation. I try to be realistic about that and I want to state this belief upfront. As well, I want to reiterate that I like all types of daylilies, every flower form/type, all colors/patterns, all heights and both ploidy levels. I think evergreens and semi-evergreens have there place, but that place is not necessarily in my garden.

So, with all of that said, on to the trait sets. Below is a bullet point list of the main trait sets that I will be focusing on for this series of blog posts.

• Disease tolerance vs disease susceptibility
• Drought tolerance vs drought susceptibility
• Fast increase vs slow increase/non-increase
• Plants that thrive in diverse conditions vs poor performers
• Healthy foliage vs problem foliage
• Dormant foliage vs evergreen foliage
• Scape strengths vs scape weaknesses
• Flower pigment fastness vs flower pigment spottiness
• Reliable flower opening vs poor flower opening
• Fertility vs infertility

I will now talk a bit about each of the trait sets to provide an introduction to my general thoughts on them.

Disease tolerance vs disease susceptibility


Disease is a given in all organisms. Nature has provided an array of methods to fight off attack by various pests and pathogens. In domestic breeding we wish to incorporate as many methods of defense into our lines as possible. However, this is no easy task. To begin with, you actually have to see the problem you wish to breed tolerance for in your garden and then you have to select those individuals that show some level of tolerance as breeders. Gardens that suffer rust, rot, spring sickness, thrip damage, or any new problems that may arise, are in a position to select toward tolerance. Gardens that do not experience a given problem are not suited to selection for tolerance levels. Yet in time diseases may spread, and if so, tolerance will become an important aspect of breeding programs in gardens that once did not have to consider such things.

An example of this in my garden is rust. I have never seen rust in my own garden. I have, however, brought in several cultivars and seedlings that have shown high rust tolerance. I will test them out for my important criteria and then utilize any that pass in my own breeding program. However, unless I start seeing rust in my own garden (which I suspect is inevitable due to increasingly warming climates), I cannot select for rust tolerance. Until such a time as I do have rust in my garden, my path will be to send select seedlings descending from these rust tolerant individuals to gardens infected by rust to see if any of the tolerance of their ancestor(s) has passed on to them. That is all that I can do in this regard at this time, but it is still a consideration for my program.

Tolerance to disease makes a given cultivar easier to grow when such a disease is present, and more attractive in such settings. Such plants are very desirable when disease is present. Intolerant plants are unattractive, may even die and at the very least present an unattractive garden display. Such plants are undesirable in diseased settings. Further, such plants are not enjoyable and will likely reduce the desirability of the daylily, in general, to the gardening public at large.


Drought tolerance vs drought susceptibility


This year has certainly shown the need to select for drought and heat tolerance. Those cultivars that are drought susceptible are unlikely to survive this year without extensive watering, and that is something I am unwilling to do, as I practice xeriscaping, being located on a southern slope with a sand and clay mix soil and I don't want to spend all my time watering. I have grown various daylilies in this setting for the last twenty-one years and many flourish, while some do not. By using those which flourish in a dry setting, I can select toward tolerance to dry conditions. Anyone with a dry garden setting can make this type of selection, if they don't give supplemental watering, or only sparingly.

With that said, I don't blame anyone who does water. Please don't think I do! There are situations where you will want to water and not everyone will want to practice xeriscaping as I do. For instance, if you have a display garden, special seedlings you wish to increase, are raising divisions to resale or have very special seeds you wish to ensure reach maturity you will want to give supplemental watering, but I would point out that watering plants in dry conditions means you may not be able to select for tolerance to dry conditions. Consistent watering in dry conditions creates a negative selection pressure wherein those without drought tolerance (susceptibility) survive and pass their genes on to future generations. If climate conditions continue to change, we will likely see more heat and drought as a result. It is something to consider. The key is to test all potential breeders in non-watering situations. I will have more to say about this in a future post.

On the opposite side of drought tolerance is tolerance to high water/humidity levels. In rainy seasons, my soil is very saturated due to the clay content and holds a great deal of water. Daylilies are remarkable plants, in that many of them can survive a wide range of conditions. There is some suggestion that high humidity/water and heat may contribute to rot. Those cultivars that can flourish in both very wet and very dry conditions are very special. I am paying particular attention to those that show such diversity of tolerance in my own program.


Fast increase vs slow increase/non-increase


The increase of fans shows a wide range of expression. Some cultivars increase rapidly, while others never seem to increase at all. The worst end of this spectrum is those that not only don't increase, but actually decrease. I have to think that most of the slow growing cultivars probably grow well enough in their home gardens, but likely that is under the most ideal of conditions. In an average garden, some cultivars just don't grow well.

I have spoken with breeders who do not like fast growing cultivars because they say they have to be divided too often. I have also heard slow growing cultivars referred to as being preferable, "hybridizer's plants", because they do not increase as quickly and do not need frequent division. My feelings are completely different. I do not have a problem with moderately increasing cultivars, but I do not like slow growing cultivars that take years to increase a few fans or never increase at all, in my setting. To me, the advantages of fast increasing cultivars is that they can be multiplied more quickly both for distribution and for use in garden settings. I also just like a really large, mature clump of daylilies.

The one disadvantage to some cultivars when they grow into large clumps is that scape count can be suppressed. However, this does not occur in all cultivars. I have grown many older cultivars for years that can be very productive and produce a wonderful display with lots of scapes when in a large, mature clump. The key to produce such plants is, of course, to use such plants as breeders.


Plants that thrive in diverse conditions vs poor performers


Some plants perform wonderfully all over the country, in every zone. These are what I call 'generalists', being that they are adaptable to a wide range of conditions. Other cultivars only seem to prosper in very narrow bands of the continent. If you happen to live in that small region where such plants flourish, they are great, but if you don't then such plants are a great disappointment when the attempt is made to grown them. While daylily enthusiasts may understand this, the general gardening public will not and can be easily turned of by highly-touted but poorly-performing, regionally specific cultivars.

I pay special attention to plants that are known to flourish all over the country. The Lenington Award winners are often good examples of such generalist plants and are a good place to start when selecting breeding stock for diverse conditions. Those who wish to breed plants that can be successful in a wide range of environments should look for plants that are known to thrive in a wide range of regions and conditions.

One of the major agricultural philosophies that I use is the Law of the Minimum. Stated simply, the necessary element for survival that is in the least supply limits the population. Daylily cultivars that survive the best in severe circumstances and that perform well in the widest range of areas and drastically different environments push the boundaries of the Law of the Minimum. By breeding from such plants, it may be possible to increase the definition of 'the minimum' for daylilies, in many regards.


Healthy foliage vs problem foliage


I pay a lot of attention to foliage. Stated simply, I detest ugly foliage, and lots of daylilies seem to have really ugly foliage. However, not all, as some show beautiful foliage, even in poor conditions. There are many types of problem foliage, but the biggest problem to me is yellowing, dying foliage. Some daylilies though will have foliage more yellowish in tone, but that survives well and does not brown and look hideous after two or three hot or dry days. While it is not my favorite color of foliage, I do not mind such plants. The foliage that I find most attractive is the blue-green foliage that does not yellow out badly and that shows good drought tolerance. There are many other considerations with foliage such as frost tolerance, heat tolerance, drought tolerance, etc. I will touch on this subject in more detail in future posts.


Dormant foliage vs evergreen foliage


Stated very simply, evergreen and semi-evergreen foliage drives me crazy in the cold parts of the year. I like those dormants best that go completely underground with the beginning of cold weather and don't come up too early. They appear clean and neat to me during the winter. There are evergreens and many semi-evergreens that survive and thrive in the winter, but they are simply unattractive to me. For anyone who they don't bother, they are fine, if they are winter hardy. For those in the south, I think evergreen and semi-evergreen cultivars are preferable. However, it should not be mistaken that I am saying that all evergreen and semi-evergreen are not cold hardy or that all dormants are only for the north. Some dormants flourish in the warm south, just as some evergreen and semi-evergreen flourish in the cold north. My stance on the foliage types is purely my own aesthetic. In spite of that, there are instances where I will continue to grow and breed with both evergreen and semi-evergreen cultivars. I will touch more on this in later posts.


Scape strengths vs scape weaknesses


Here is something of a major pet peeve to me. I simply hate weak scapes, in most instances. I prefer scapes that are perfectly vertical. I can tolerate scapes that may lean out as much as 15-25 degrees. Any lean beyond that is a major flaw, in my opinion, no matter what the flower looks like and makes me want to compost the plant immediately. I might on rare occasion use such a plant as a breeder if there are other very exceptional performance qualities, but that is a great rarity. The only exception to this rule is with very short border-type cultivars such as Stella De Oro and descendants where the trait is not horrible, but I don't really even like it in that group of cultivars.

There are several types of scape weakness. Some just fall over at the base or the base of the scape cracks or bursts. Other scapes are strong at the base, but then get thin part way up and curve over. Some scapes simply can't hold up their large flowers. Large flowers need very thick, strong scapes. Some cultivars have very thin scapes that haven't the ability to hold up any sized flower.

I have had people tell me that a given flower is so wonderful it is worth staking, but not in my garden. I have better things to do than run around staking up weak stems, especially when there are so many lovely daylilies with strong scapes. I will have much more to say about this trait in future posts.


Flower pigment fastness vs flower pigment spottiness


Again, a really simple thing. I don't like spotty, splotched, melted flowers, even in heavy rain or bright, strong sun. There are cultivars from every color that do not show spots and pigment bleed/melt. I see no point in working with flowers showing these flaws, as if there is any genetic basis to these traits (and there likely is), then using such plants probably passes the problems into future generations. More to come...


Reliable flower opening vs poor flower opening


I will eventually have a lot to say about this, but in simplest terms, flowers that don't open well are not popular with me. Twisted messes, stuck pie crusts, faded blooms gumming up other blooms, trumpet forms on cultivars that are supposed to open out fully, flowers too close together to allow full opening; these are things that make me contemplate the compost heap. I am more understanding of poor opening on late fall rebloom or on very cold morning. What I really hate is when the plant in optimum conditions and its normal season of bloom still doesn't open well. We will explore this more in future posts.


Fertility vs infertility

Fertility is very important to me, not just for my momentary gratification or to get out as many seeds as possible (though that is not a bad reason in and of itself), but because our selection of breeders influences the future of our lines. Using breeders with fertility problems may cause fertility problems in future generations. The issues of fertility in daylilies is a big subject and will require future posts to be explored well. In short, I place strong emphasis on breeders that show strong pod fertility with preference for strong pod and pollen fertility in a given cultivar. I may at times use pollen parents that show little or no pod fertility, but they should then always be crossed over strongly pod fertile cultivars and any breeders chosen from such a cross would be backcrossed to strongly pod fertile plants with further selection toward pod fertility applied to subsequent generations. The ideal cross though is strongly pod and pollen fertile x strongly pod and pollen fertile, where possible in my own breeding program.

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In future posts, I will write about each of the above criteria individually, looking at them in greater detail and giving more of my opinions and observations on each. I hope you enjoy these posts and find them useful, but please always remember that these are only my thoughts, opinions and observations and are only meant to apply to my own garden and breeding program. You may have different criteria and some of mine may not apply to you at all. In such a case, do not take it that I am saying my criteria should apply to you. Thank you!!

Brian Reeder

zone 6/7

AHS region 10