Wednesday, January 17, 2018

The Daylily as a Permaculture Subject - Part 3

The Daylily as a Permaculture Subject 

Part 3
Breeding for Improvement 

Brian Reeder

*Disclaimer - (I am not recommending in any way, form or fashion that anyone should consume any part of the daylily. While Hemerocallis (daylily) is a common food and medical substance in Asian countries, if you have never eaten daylily or do not have accurate information on the consumption of daylily, it is completely upon your discretion when experimenting with consuming daylily. I strongly recommend that first time consumers be very careful in the amount of any given daylily they eat in order to observe the effects on their individual systems. Some people have experienced diarrhea from consuming daylily in quantity and some daylilies have been known to have sedative effects, which might be referred to as ‘poisoning’ or ‘poisonous effects’ by some sensitive people. The internal chemistry of each daylily, as well as each person, may vary and so no general information about daylilies as food or medicine can be made that will cover all daylilies and all human reactions/interactions with/to daylilies in any given instance. What is perfectly fine for one person may not be for another.)

In the final installment of this series we will look at the possibilities of forming a breeding program to select for traits in daylilies that can increase their value to permaculture, both in terms of desirable plant traits and nutraceutical traits.

I have mentioned several times in the past two articles that I have been making some selection in this direction, with a focus on plant traits that make for a better garden as well as permaculture specimen and flower traits that make for better display uses as well as culinary applications. This does not mean that any of these plants are separate from flower trait selection (in terms of floral display) in my overall floral daylily program. The traits that make for a better permaculture specimen also apply to making a better and more garden-worthy floral specimen. As well, one of the most desirable traits in a good culinary daylily flower is also of utmost importance to a good garden specimen of floral display daylilies - i.e., resistance to thrips, aphids and other insect predators that cause damage to or destroy the flowers, buds and sometimes even scapes.

A hybrid daylily with high susceptibility to thrips, showing extreme damage to buds and scapes, and foliage heavily damaged from late freezes. I suspect that the combination of late freeze susceptibility and thrip susceptibility can combine to make a much more severe effect. You can see the beautiful, deep green foliage of the plant in the background to the right, which has strong late-freeze resistance and is a later blooming cultivar and so showed no damage. 

So my permaculture selection isn’t a separate part of my program, but rather, is composed of plants from within my overall selection program that are attractive, vigorous, hardy and show high resistance to disease and insect pests and just happen to also have very good tasting buds/flowers that have no negative effects on me physically when I eat them. So while there may be people out there who’s only interest in daylilies would be to breed for permaculture applications (and may select for a specific type of flower in conjunction with their program, or not), anyone who is breeding daylilies for any reason, who pays any attention to plant traits, could theoretically be selecting plants that also show high culinary value (though I am not suggesting everyone should be selecting for culinary or permaculture traits, either). However, such selection does not have to be a ‘separate thing’ from daylily flower breeding, though it can be. That depends on the interests of the breeder. I just want to stress that this does not have to be a separate thing, a whole world unto itself, unless the breeder wants it to be.

The crosses that have shown the greatest application for culinary uses in my program, and that are also giving me the best range of desired plant and flower traits, are the base crosses I have made to build my future floral daylily program upon. In fact, the plants that are showing the best applications for permaculture purposes in my program happen to be a side effect of the outcrossing I have done between modern hybrids over species and species-like material - specifically seedlings coming from Hemerocallis fulva Korean, Hemerocallis Hankow, Notify Ground Crew and Ancient Elf.
So this is how my own breeding for permacultural uses emerged - by chance as a secondary side effect of my program to build a new tetraploid base-stock by going back to the species and species-like hybrids to open the tetraploid gene pool. The selections I have made are selections I would have been making anyhow, and my focus has been a vigorous, hardy plant with high disease resistance and a lovely flower with high pest resistance. By chance some of those selections have also turned out to be delicious, but if one wanted to make actual crosses and selections toward specific goals of permacultural or nutraceutical uses, that would be possible as well. In fact, I am now beginning to make some crosses of this nature, now that I have plants that I feel would have special application for these purposes.

So what could one be selecting for? On the most basic level, one could be making crosses and selection for the most basic and desirable traits - hardiness, vigor and disease resistance of the plant and flowers of good flavor with high pest resistance. However, there are probably many other traits that could end up being bred and selected for. Since (apparently) no one in the west is actively breeding and selecting for nutraceutical traits, it looks to me as if this would be an open field, where almost any type of program could be ‘unique’ or nearly so.

In terms of food applications, some daylily flowers taste very sweet, almost like very sweet lettuce, while others are more bitter, less sweet or more astringent in flavor. Some say that one color or another is sweet and some others are bitter, etc., but I have not found this to be the case, and I cannot say to you that a given color is “good” while another is not. I find it is much more variable - one yellow is delicious, another is less so, one purple is bitter while another is sweet, etc. The species and near-species hybrids that I have previously discussed all taste good to me, for instance. 

Some daylilies have been shown to have high protein levels and high vitamin C levels in their flowers, but I don’t know of any testing across many types to determine variations. This may be a open field for selection. Other flower traits that might be selected for, combined with flavors and nutritional content, would be flower size, texture and storage ability. A delicious, nutritious and large flower could be very enticing, especially combined with high pest resistance on a vigorous plant with disease resistance, hardiness and many buds on the scapes. Let your imagination be your guide and experiment with what is possible. I expect a great deal is possible once attention is paid to the underlying trait variations.

Whooperee - An exceptionally thrip and rust resistant evergreen cultivar.

In experimenting with eating daylily flowers I have also run across some interesting effects, in addition to flavors. As mentioned in the first part of this series, daylilies have traditionally been attributed with medicinal properties, especially sedative effects, and modern research (as noted in that first part) is also beginning to demonstrate that this may well be the case. In my own experience, this is the case, and the effect(s) and strength varies from one plant to the next. Some plants have little to no effects at all, and these are the most desirable, in my opinion, as a food source. Others will have a mild sedative effect that manifests for me as a slight haziness that is relaxing but does not interfere with my ability to function. 

Still others have a considerably stronger effect and these can be so strong as to be described as strongly sedative and even mildly hallucinogenic, creating a euphoria combined with visual lights and colors, especially when the eyes are closed, and stimulate very deep and relaxing sleep, with no notable effect upon waking the next day. It is my experience that these are quite rare and are probably responsible for the reports of daylilies that are ‘toxic’ or ‘poisonous’. However, I have not experimented with any of these types enough to give any definitive report and I do not recommend you try any daylily flowers except in very small doses (one petal, perhaps?) until you find its effect on you, as well as your body’s tolerance to daylily flowers in general. If you see no negative effects, you might increase the amount you eat at a given time to see if there is a dosage effect in larger quantities.

The strongest effects I have experienced have been from eating five or more buds/flowers (and fresh buds seem to have stronger effects than dried and cooked flower) so perhaps some of the ones with milder effects would show stronger effects in larger quantities. However, since these pharmaceutical effects do not hold much interest for me, I have not experimented very much with these effects and so can give very little information and can offer no recommendations, except to say, that for those who might have an interest in these effects, whether permaculturalists, medicinal/pharmaceutical researchers or interested hobbyists, there is seemingly variation and undoubtedly these are areas that can be modified (perhaps highly so) through selection. Good research in labs by professional researchers would be of much value, both for pharmaceutical effects and nutritional factors. Without access to such testing, hobbyists and permaculturalists can still make progress on selection for such traits, though I strongly recommend they proceed with caution, especially in terms of the pharmaceutical applications.

Further, as the research cited in the first part suggests, there may be even more pharmaceutical effects in Hemerocallis and these traits may well be amenable to selection as well. However, I feel these may be more difficult to select for, unless one has access to laboratory testing. I personally wouldn’t have a clear idea of how to do such selection without access to chemical and perhaps genetic testing, as well as application trials in non-human test subjects. I think this work is beyond the scope of most hobbyists, but may be applicable for some permaculturalists, especially where that person(s) is a trained researcher with access to lab testing. I want to stress that I am not suggesting anyone should attempt to do work with traits of a chemical nature that they are not qualified to work with, but these traits may well exist in the Hemerocallis, and if one has the skill-set to work with such traits, I would still hope they would apply the other breeding suggestions I am recommending herein for working those traits into strong, vigorous and disease/pest resistant lines.

The traits and uses that I want to stress here, both because I think they are the most applicable traits for hobbyists and home permaculturalists to work with and because they are the traits I am personally focused on and feel I can comment on best, are the plant traits that make a great plant for permaculture/food forests/landscape specimens and that make flowers stronger, durable, resistant to pests and useful as a floral or food item. I feel these are easy traits to work with and can have the greatest application. Both are complementary for those already breeding daylilies for flowers and both will be of value to those who solely have an interest in daylilies for permaculture applications. 

(Gram's Dream x Arctic Snow)
A seedling from my program showing the type of plant, scape, bud count and flower I like - strong, hardy, healthy, disease resistant, brightly colored and unblemished.

With that said, I do want to stress that I feel permaculture shouldn’t eschew beauty and can engage in any level of flower phenotype selection that might tickle the fancy of that permaculturalist. I do not know if all daylily phenotypes will have applications for food purposes. I haven’t eaten flowers from every daylily cultivar or seedling I grow. Some flower phenotypes may be pleiotropic with, epistatic to or linked to traits that would cause problems with food applications. However, where any flower trait can be combined with good food qualities, then the sky is the limit.

Fortune's Dearest 
A lovely purple color and white tooth edge to petals and sepals - a lovely look from modern floral daylily breeding.

I also want to stress that I am not preparing this series to create sales for myself of either daylily species or cultivars or my own introductions. My goal is to inspire others to look at and experiment with the daylily beyond those who are already floral daylily hobbyists, and with floral hobbyists, beyond the singular focus on flower appearance. I do sell H. fulva Korean, Notify Ground Crew and Ancient Elf, but my stock is small and I have no interest in becoming a large commercial seller, nor do I have any interest in selling out of these. I do want to promote these plants and I want to see many people growing and breeding from these and other good cultivars, but whether they are sourced from me or another vendor is irrelevant. What is relevant is to spread the word about what I am finding that is working best for me and for other breeders whom I communicate with. I hope to see many other growers and vendors offering these, regardless of where they obtain their initial stock for propagation.

Further, I will be introducing plants in time that I consider highly applicable to permacultural uses. In all instance, where I know the parentage (which will be most of the time), I will list that parentage. In that way, you can buy these cultivars from me (and I do hope to spread my best work around for others to grow and use), but more importantly, I want to get the information out to others as to what I have worked with and what has given me the best results so that others can work within those same genomes and use them to make their own unique combinations and selections. Further, my best selections will not always fit every environment, so others need to be tailoring this type of breeding for their own environments, even if they do work with some of my introductions. 

(H. fulva Korean x Queen's Circle)
A seedling and future introduction from my program that has many applications for both floral and permaculture breeding. 

My goal is to increase knowledge and interest, as well as to inspire others to also pursue work beyond the increasing and intensification of the flower traits. If you want to buy the best results of my breeding, testing and selection, I certainly will not complain, and I understand that some people will not want to take on a breeding and testing program of the scale I have and so may prefer to buy improved stock rather than breed it up themselves. However, for those who wish to do the breeding themselves, take the information I offer and run with it! Take it in directions I have tried and in directions I have not tried. Put your own unique stamp on it and select it to your specific garden needs and your own tastes and aesthetic. 

(Notify Ground Crew x Arctic Snow)
A seedling and possible future introduction from my program that has many applications for both floral and permaculture breeding.

More than anything, I want to inspire people who might not really think much of daylilies to begin thinking about them and growing them. They have many uses beyond flower gardens. Since there is little available information about these other uses, it is my hope to spread some of what I have found through trial and error and to offer you useable information to make good selections. The greatest asset of the hybrid Hemerocallis is that there are a great many varieties and they show great diversity. However, this very thing can also be their greatest drawback, as they are not all created equally and making good selections for uses beyond the flower can be extremely difficult due to the lack of reliable information. I hope to help make that information a little more accessible and to inspire you to seriously consider the daylily.

Now I want to touch on my experiences and offer an assessment on a few of the traits I feel are important to permaculture uses (and floral daylily breeding, as well).

1. Rhizomatous Spreading - Usually said to be recessive in the floral daylily community, but my observation is that the trait appears to be a single major dominant gene with simple Mendelian behavior. There are likely modifiers as well, but the appearance is of a single gene that is dominant with a dosage effect. Homozygotes show greater spreading behavior (thought this may vary in just how much, how far and how vigorously - thus the possibility <likelihood> for modifiers). Heterozygotes show spreading behavior, though less so, spreading slower and each fan arising at a shorter distance from the main plant. All the fulva clones I have grown are rhizomatous to some extent or another. Some daylily hybrid cultivars show rhizomatous behavior such as Bill Fall and La Vida Loca. The trait is generally considered undesirable by floral daylily hobbyists, but can have many applications for landscaping uses and permaculture uses.

A patch of rhizomatous daylilies

Hybrid tetraploid daylily cultivar Bill Fall

H. fulva Korean growing in large containers. This is an excellent method to restrain rambunctious rhizomatous types of daylilies. Some running types of daylilies can escape and become wild, invading surrounding areas, so care should be taken when growing such types if their escape is a concern for you. Containers allow you to maintain such plants without fear of escape. Just keep an eye on any runners that try to emerge from the holes in the container to be sure they don't establish in the surrounding soil. Not all running types are invasive enough to be a concern, with H, fulva ex Europa, commonly called the ditch lily and ubiquitous throughout much of the world, being the most extremely invasive type.

2. Clump Forming - While it is generally thought of as dominant in the floral daylily community, clump forming seems to be recessive to rhizomatous growth, in my experience. All of the yellow species are clump forming and most hybrid daylilies are clump forming. In tame flower gardens, clumping is generally preferred, but in many permaculture applications, the spreading habit of the rhizomatous types may actually work better.

A beautiful display from the clump-forming daylily cultivar Spider Man. Spider Man is a fairly dormant plant that shows high thrip and rust resistance in my experience.

3. Rust Resistance - Seems to involve multiple genes, both dominant and recessive. Daylily plants that show very high rust resistance in field tests can often show very good breeding value for rust resistance. Because of the multigenic nature of rust resistance, I can’t tell you any percentages to expect, as that will vary wildly. The genetics of rust resistance is complex. However, I can recommend that H. fulva Korean and Ancient Elf have shown exceptional rust resistance and breeding value for rust resistance in my own program. To test daylilies for rust resistance, you must expose them to daylily rust and you must not spray them with fungicides to combat rust. To breed for rust resistance, it is best to use the most resistant plants you can find as parents, and their seedlings must be exposed to rust. Without rust, you can’t really select for resistance to rust. However, even if you can’t select for rust resistance, you can at least base your lines on plants that show exceptional rust resistance.

A susceptible plant to daylily rust (behind) showing heavy infestation with a very highly resistant daylily plant (foreground) showing no rust infection at all.

4. Thrip/Pest Resistance - While I have seen daylilies that appear to be fully immune to rust, I have never seen any daylilies that seem to be totally immune to thrips. However, I have seen a few cultivars and seedlings that show high thrip resistance. I have been breeding and selecting for thrip/pest resistance and it seems that those plants that show very high resistance to thrips are able to produce seedlings that also show very high resistance. I cannot yet say that this trait is a single gene or multigenic, or that it is clearly dominant or recessive. What I can say is that it is apparently heritable. I will need to observe for some years more and with a good number more crosses to have a good feel for what to expect in breeding and selecting. What I can tell you now is that the trait(s) appears to be heritable when thrip/pest resistant plants are used as parents. The most thrip resistant plants I have found include Solaris Symmetry, Whooperee, Spider Man, Notify Ground Crew, Tis Midnight and Corky (and its descendants including From Darkness Comes Light, etc.) As with rust resistance, you need to expose daylilies to thrips and other insect pests to select for resistance and you must not spray pesticides to eliminate these pests.

Hybrid Daylily cultivar Solaris Symmetry

5. Flower Flavor - I have only been selecting for this trait for a couple of years and haven’t seen enough seedlings to make a guess as to how many genes may be involved or how those genes may work, but I can say that the trait seems to be heritable. In other words, daylilies that taste good to me have produced seedlings that taste as good or nearly so, though sometimes they also produce daylilies that have flowers that do not taste good. All I can tell you at this time is that there seems to be heritability for the flavor traits. I will address this more in the future as I have a better idea of what is happening, but I wanted to tell you that there is apparently room here for selection, so please, do try if you are interested.

6. Flower Color (Visual Anthocyanin versus Non-Visual Anthocyanin) - I want to address this in the most general of terms. This is actually a very complex area, but at the most basic level, we can look at the two classifications the Chinese use ( ‘yellow’ vs ‘orange’). For me, ‘yellow’ encompasses all the non-visually anthocyanic types such as all tones of yellow (carotene) and all tones of ‘melon’ (a seemingly-recessive gene that changes carotene to lycopene), pigments that are deep within the tissue of the flower petals and are also present in the visible-anthocyanic group in the petals underneath the anthocyanin serving as ‘base-color’ that can effect the visual tones of the anthocyanin as perceived by our eyes. 

The ‘orange’ group includes all those that have visible anthocyanin (a water-soluble pigment in the upper level of the petal) ranging from orange to red to purple to blackish-red to blackish-purple and also including all pink and lavender tones and possibly some near white flowers (near white is likely occurring on both anthocyanic and non-anthocyanic types). I also want to mention that I refer to “non-visual anthocyanin” (the ‘yellow’ category) as such because these almost certainly contain anthocyanin that, as shown by putting these flowers under ultraviolet light, would be visible to insect eyes, but not to human eyes. Visual anthocyanin may be present on the entire flower, only on the petals (and not on the sepals or vice-versa) or as an eye, either on visually anthocyanic types or on non-visibly anthocyanic types.

I am not going into details about the heritability of all these different variations. What I want to focus on here is the most basic level. I suspect that the fulva is older, evolutionarily speaking, than the many yellow species. I suspect that non-visible anthocyanin is a simple ‘knock-out’ gene. That is, a gene that stops the visible anthocyanin from forming in the flower. My experience is that visible anthocyanin is dominant, while non-visible anthocyanin (or absence of visible anthocyanin, if you prefer) is recessive and that this is a simple, Mendelian single gene. The complexity comes after one or the other of these two pathways are active. There are other genes that determine which type of anthocyanin you get (each color of anthocyanin is likely something different genetically and are different types of anthocyanin such as delphinidin, which makes purple tones, etc.), while yet other genes are responsible for eyes, bicolor, self, dilution of tone/intensification of tone, etc. 

An example of a yellow daylily (left), showing no visible anthocyanin, (center) under ultraviolet light where patterning that may be anthocyanic in nature is revealed. Picture right is the same flower in grayscale.

My experience is that at its most basic level you either have visual anthocyanin (which is dominant) or you don’t have visual anthocyanin (which is recessive). This actually makes breeding fairly easy. At this most basic form, let’s do one example - yellow and red (of whatever shade or tone). A red, having visible anthocyanin which is dominant, can have one or two doses (at diploid level) and still be visually red (of whatever shade or tone). When crossed to the yellow flower without visual anthocyanin (no eyes, etc., just yellow self), the homozygous-for-visible-anthocyanin-red creates 100% of its seedlings that show visible anthocyanin (red, orange, reddish tones, red or orange bicolor, red or orange eye, etc., depending on the other modifier genes present). When the yellow is bred to a red that is heterozygous for visible anthocyanin, you will see 50% seedlings showing visible anthocyanin and 50% of seedlings that do not show visible anthocyanin. Some of the 50% showing visible anthocyanin may just be an eye or bicolor, etc., depending on the other genes present. Tones of each may vary, but the split at this basic level seems to be quite reliable across a huge number of crosses I have made.

Crosses between different tones/colors of visibly anthocyanic flowers or between different tones of non-visibly anthocyanic flowers are not simple and can produce a wild range of tones. This indicates to me that there are multiple genes involved in the exact color and tone of visible anthocyanin and the non-visibly anthocyanic, as well. Neither group seems to be straightforward or simple in terms of exact colors, combinations and patterns and must represent multiple genes.

I hope this gives you a basic overview. I have tried to give you some suggestions for base plants, plants for rust resistance, plants for thrip resistance and some idea of what to expect (at the most basic level) in terms of breeding for some of the basic traits that may be of interest in breeding daylilies for permaculture purposes. For anyone who is interested in pursuing this type of daylily use or breeding, please feel free to contact me through my Facebook daylily page, Sun Dragon Daylilies, or through the email address found on the about/contact page at my website Sun Dragon Daylilies. There is much more to the daylily than just the flower and I hope a new generation of breeders will pursue these traits in addition to the beauty of the flower.

Part 1     Part 2