Thursday, October 31, 2013

Some Halloween Horror Stories for the Daylily Breeder

Scary Stories for Daylily Breeders

In my many years of gardening, I have been fortunate to be friends with a great many prominent Master Gardeners, Plants Breeders, Landscape Architects and Designers. I have remained friends with many of these people for the last few decades. They are interesting people. Interestingly, I know many of them socially, not through gardening, but through mutual friends, though some I know through various garden interests.

Since I began seriously breeding daylilies (as opposed to just growing and collecting them) in 2009, I have been speaking with these friends and acquaintances about their views on daylilies. They have had some interesting and often unflattering things to say. I take their opinions seriously due to their prominent positions in their fields, but also due to my respect for them and their work.

As I have always had a focus on the breeding of disease resistance in both plants and animals, this is a subject I always get around to in my discussions with these friends. In the last couple of years, I have been making rust resistance an important selection point in my daylily breeding and so that has often been a matter of discussion with my friends. They have strong opinions. Over the last year, I have come up with a list of very pointed and specific questions that I have been asking these people in regards to daylilies. 

Before I get into the feedback I have been receiving, I want to also mention that I have been building relationships in the daylily world over the last few years, and it is common knowledge behind the scenes that daylily sales have been in decline for about a decade now. Many prominent breeders have told me this is mostly due to rust, while a few also mention the economic downturn of the last years and some point out that people have become tired of the exorbitant prices of new introductions, especially when so many aren't that different or have problems. One told me (and some of you reading this will know who this is by having been told the same thing by this well-known person), "The golden age of daylilies is over." 

So we have good evidence that the interest in daylilies has been declining in the daylily world itself, and while we may debate the causes (and there are likely several), rust has in no way helped. With this in mind, I wanted to find out what the rest of the gardening world was thinking about daylilies and if that same decline was being seen in the non-daylily-specialty gardening world. Unfortunately I have to report that it is.

When in conversation with my gardening friends, many will mention that they have mixed feelings about daylilies. Some feel the foliage is ugly and thus the plant is only good when in flower, which outside the few reliable rebloomers, is about a month out of twelve months in the year. However, in spite of this, some of my friends still use daylilies and simply find ways to mask the foliage when the plants are not in bloom. One clever friend plants daylilies with tall scapes behind small shrubs and medium sized perennials to block the foliage, but still see the flowers.

Many of my friends prefer older daylilies that are well tested and reliable for garden use. While they generally like the flowers of the newer, ornately flowered cultivars, they find them useless in the landscape for many reasons, including the large wilted flowers needing to be daily removed, the edges hanging and the flowers not properly opening, the flowers being overdone and not working well with a landscape design, and (the one I hear the most) the poor performance of many newer daylilies compared to the older cultivars.

One friend told me, "Daylilies had always been so hardy, but they were very plain, then they bred up nice flowers and nice hardy plants and I still use some of those, mostly from the 1960's through 1980's, but as soon as they did that, they immediately started breeding those overdone flowers with poor plants that don't do well in the garden and don't work well in landscape design and ruined the daylily as a garden plant." Another said, "Why don't daylily breeders work on nice foliage? The foliage is visible much longer than the month of flowers on most of them. They are often wonderfully hardy, but I just don't get how in 100 years, they couldn't have improved the foliage."

However, the responses in regards to rust have been both the most interesting and most disheartening. Only two out of 17 of these people have not encountered rust to some level in their work. Of those 17, 12 will no longer consider daylilies in their work or gardens or will only use older, tried-and-true cultivars that they have found to have suitable resistance. None of them are willing to implement an expensive spraying program for daylilies. Of the five who will still keep or use daylilies, two are those that have not yet had rust. 

On using daylilies in landscapes and gardens
This one gets a lot of replies, often quite heated. Three of these people didn't much care for or use daylilies to begin with. Several are just adamant they won't use daylilies anymore, but did before rust swept through various plantings. Some have kept the cultivars in their gardens and customer's gardens that showed little or no rust and divided those, using them in place of the susceptible that they threw out. Some just threw them all out.

One friend said, "I loved using daylilies, especially the older cultivars with good scape height that could have the ugly, yellowing foliage hidden but the flowers still seen, and those older cultivars had simpler, more wildflower-like flowers that worked best in my landscape designs, but since rust appeared, I am much more careful about using daylilies in my clients landscapes." He recounted to me a horror story of a client who was fond of daylilies and so they had used a good number of newer (read expensive) cultivars in her landscape design only to have most of them rust over the first time rust entered the area. He offered to spray the daylilies for her, but her reply was that she really loved roses, but didn't keep them because of the spraying they required and she had only been growing daylilies as a substitute for roses, so she certainly wasn't going to be spraying daylilies! They ripped them all out and threw them in the trash.

One friend, who is a well known and very in-demand landscape designer and Master Gardener had a lot of things to say that I can't repeat in a public forum due to all the four-letter words, but I will distill the essence here for you. He feels that the daylily world is directly to blame for not getting on the ball and being more honest and forthcoming about rust, and especially for not making a concerted effort to identify resistant cultivars and to then breed up more resistant cultivars. He says that every successful garden plant has at one time or another faced disease issues and that it is the responsibility of the breeders to breed for resistance. He says that anyone breeding anything should know from numerous examples amongst other plant types that resistance breeding is possible. He says that he will never use daylilies in any of his projects again because you can't reliably get information on resistance and that the breeders seem not to care at all about their clients needs and he won't forget it. (I did go on to tell him about several cultivars I know to have reliable resistance in many locations and he says he will consider trying a few of those, just to see what happens, in his home garden. I will keep on him and see if he can't be convinced by a few of these. I just sent him a box of plants that have been reliably resistant here, for him to try out and I have my fingers crossed, and you should too, because this is someone with a huge platform and a powerful voice in the plant and landscape world!)

On spraying daylilies to control rust -

This question has prompted some interesting reactions. The biggest reaction is peels of uproarious laughter. The other most common reaction involves lots of four-letter words that are unprintable here. In short, none of these people are remotely willing to start a spraying program for daylilies.

One friend said, "Why on earth would I waste a clients money to spray a disease-prone weed?" That was the kindest thing he had to say.

Another said basically the same thing, adding, " The daylilies have always been little more than wildflowers in my landscaping work and they are not worth the time, bother and expense of spraying when I can just rip them out and put some of the wonderful new coneflower cultivars in their place."

Another said that if he were going to start an expensive spraying program that had to be repeated throughout the year, he would rather start using roses. He said there was no chance of him advising a client to plant daylilies and then take on the expense of a spraying program.

In all, none of these seventeen people saw a spraying program for daylilies as a wise investment of resources and effort and none said they would even consider it. The consensus was that they would either stop using, advising and growing daylilies altogether if spraying is a necessity or that they would only use/grow those that did not require spraying. The overall view was that daylilies did not have enough value in their gardens or design work to merit spraying, and one went out of his way to point out that daylily breeders needed to realize this fact!

On rust resistant daylilies-
This question drew, for me, the most interesting and promising responses. There are two main points or lines of questioning here. One is - "Would you use daylilies if you could get rust resistant cultivars?" and the other is "How resistant do daylilies have to be for you to feel they are usable, or said another way, how much rust can you tolerate?" 

On the first question, in all instances except the three people who didn't like daylilies to begin with, all the respondents said they would be interested in using daylilies again or beginning to use more daylilies in their projects if reliably resistant plants were easily obtainable. They all felt these didn't have to be new resistant cultivars, but could be any of the older cultivars (no matter how old) that showed good to excellent resistance. All felt that resistant cultivars were the only solution to rust in the garden because spraying was not a reasonable or viable option. Several had already been using those cultivars they had or that were in their clients gardens that had shown moderate to no rust, and some had torn out the rust magnets to replace them with divisions from those more reliable cultivars.

On the second question, none of them felt that total immunity was required (even the three who didn't like daylilies), but that moderate to high resistance was desirable and would be enough. One said, "I don't mind a little rust and my clients don't see it if it doesn't get out of hand. The real problem is those cultivars that are simply covered in rust, because they are wretched looking in the landscape, my clients notice them and complain and their performance is often compromised in the following season, if I haven't already removed them." Another said, "The problem is not a bit of rust and absolute immunity may be unrealistic in any species. The problem is those that look like fountains of orange felt spraying to the four winds. These are the plants giving such a bad name to daylilies, even those that don't show it as badly." A third said, "If my clients don't notice it, and it doesn't kill the plant, then there is no problem." 

I believe this applies to average gardeners too. For instance, when I first showed my mother and my aunt a rusty plant, they couldn't tell there was a problem and I had to bring up a leaf and show them the rust on it for them to see the problem. They both commented that, "...daylily leaves are so ugly and yellowed by this time of the year anyhow, I just thought that was all that was." Most gardeners are going to feel the same and not be bothered by a small amount of rust. However, the full-blown rust magnet is a different story. My mother and aunt can spot those across the garden and they don't like those and I end up getting to pull those and add them to the trash bin. This is basically the exact sentiment I keep hearing from my professional gardening friends.

What I found amongst these professional people is that they have an understanding that total immunity is unrealistic, that any pathogen may mutate and thus breach the resistance of a given plant and that all resistant plants may not remain so in perpetuity. One said that he uses lots of coneflowers (echinacea), especially the newer cultivars, but that they don't last forever and often are replaced with seedlings of lesser merit, so he rips them out and replaces them every few years, and he wouldn't feel bad about doing that with a daylily either. He felt that if he got three, five or even ten years of good use out of a cultivar and then it failed in resistance, and could be replaced with a newer cultivar with resistance, even if again it only lasted of a number of years, that this would be no problem. As he says, "The perennial plants in a garden are generally not permanent installations." What he is troubled by is purchasing a daylily, planting it and then having to pull it out after one season because it became a rust fountain. He says that daylilies are simply too expensive to be treated as annuals...I can see the logic.

So in closing, I hope our tour of terror hasn't caused you too much stress or fear. Things are changing for the daylily. Rust is a pressure that isn't going away and will reform the daylily world, no matter how much we resist. If we can accept this new reality and flow with it, I think we will be much better off than if we ignore it, remain in denial and pretend that expensive, repetitious and labor-intensive spraying programs is an answer. These programs may be reasonable for sellers, but not so much for most gardeners.

So let us look again at the main thrust of the feedback I have received.
  • Aside from rust, the biggest complaint about daylilies is the less-hardy new plants with overdone flowers that require pruning of spent flowers and don't open well, as well as less-than-attractive foliage (on old and new cultivars). To me, focusing a little less on how many edges are there and a little more on how the foliage holds up could be a shift in the right direction to keep the daylily appealing beyond a small group of collectors.
  • Rust is turning landscapers, garden designers, Master Gardeners and gardeners in general off from the daylily, and may not be helping the popularity of the daylily amongst daylily people either. Most specifically, it is the very rust-susceptible (the 'rust magnet' or 'rust fountain') that are causing this feeling.
  • Landscapers, garden designers, Master Gardeners, and general hobby and home gardeners are not very interested in starting a spraying regimen against daylily rust, especially when they do not view the daylily as a top-line garden plant and have so many other hardy and spray-free choices for their gardens.
  • The Master Gardeners, landscapers, garden designers and general gardeners I have spoken with do not expect total immunity against rust. They would be content with good to high resistance manifesting as moderate to low rust sporation in their gardens and client's gardens. Most are realistic in understanding that resistance is tenuous and may not be permanent. They would be content with some resistance and would continue to utilize daylilies that show some resistance and do not become unsightly due to massive rust sporation.
I do hope readers will understand that I am not serving up this dose of fright just for the sake of inflicting fear, but to address a real problem that is slowly creeping through the daylily world and from there out to gardeners of all types. It is my intention to offer the content of my conversations for your consideration so that you, the daylily community, can begin to make some strides toward helping our much beloved friend, the daylily, remain a desirable and attractive garden plant for many years to come.
Happy Halloween!

Friday, October 25, 2013

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