Wednesday, July 22, 2015

Bridge Plants

Three Facebook Posts on Bridge Plants

I have condensed these three Facebook posts into one article.

I wanted to talk a little bit about bridge plants and use the attached picture as an example. A bridge plant is a plant that has traits (or carries traits) that you wan't, but isn't good enough to be an introduction. It is simply a breeder in your program that you are using to bring in traits you want.

This particular seedling pictured below is Ashwood Wray of Sunshine x Pigment of Imagination. It is an inbreeding I did to see what would happen and this is the only seedling I kept from the cross. It isn't the prettiest flower ever, and is definitely NOT an introduction, but it has some amazing traits. For instance, it puts up four to five series of scapes each year I have had it since its first year of flower three years ago. That is some impressive and consistent rebloom! It has been totally rust resistant as long as I have had it. It has beautiful foliage and it increases quickly. Finally, it shows the odd blue of its pollen parent as an undertone (though it is not a color changer) at the edges and eye and it is a white based flower rather than yellow based, which can give color clarity in breeding.

However, look at the stamens - they are withered in the first few sets of scapes and even in the last sets of scapes it have never produced viable pollen - it is always white and sterile. It does not set seeds on the first sets of scapes, but on the last two sets of scapes in the fall, it sets seeds like crazy. It is top branched and doesn't have a great bud count, but with the constant rebloom, it doesn't really matter that much. The color is nothing to write home about and it is a bicolor, but there is the bluish undertone, which you can see strongly at the edges and in the base of the petals just above the throat. Finally, it doesn't have a great form and certainly doesn't show the flatness of its pod parent or its ancestor Substantial Evidence.

So what to do with such a seedling? It has too many good traits to cull, but it will never be introduced, so it is a bridge plant to use in breeding. I have been using it as a pod parent in the fall on the scapes that will set seeds. I am crossing it with other reblooming line plants (mostly my own seedlings) that are totally unrelated to the Substantial Evidence family line that this seedling comes from. I am careful to use it only with very pollen and pod fertile partners and have been focusing it through deep reds and purples that show a melon/near white base. I have gotten some very vigorous seedlings from it to date and a few should bloom this year. I will be curious to see what the seedlings from it look like and if I will see the pollen problem in any of them. However, whether they do or not, I will watch any descendants of this plant closely for pollen problems because I know this one has it. I consider this a "salvage plant" (for me - often a synonym for 'bridge plant') and its use is a "salvage project". That is, I am seeking to salvage the good traits of this plant and improve or remove the bad traits.

Yesterday I looked at a situation where there was a plant with a defect that is being used as a bridge plant because of other desirable traits. Tonight, I want to look at a group of seedlings that are also bridge plants, that don't have any obvious flaws, but still do not merit introduction, though they are important breeding subjects.

The seedlings pictured with this post are from seeds given to me by Mike Huben in winter of 2011 and are from his 2010 breeding season. They are a cross of his cultivar Early and Often x (what was at that time) a seedling - MH0735X - which is now registered as Army of Darkness. Mike made the cross to combine rebloom with dark scapes and possibly also to get a pale melon to near white flower. The seeds were started here in spring of 2011 and the plants are now 4 years old.

Mike told me he was disappointed in the cross because he didn't get dark scapes, rebloom or near white flowers from any of the seedlings. My reply was that they are exceptional plants and all carry all those traits recessively, so it is just a matter of going another generation to pull those traits out. Mike has a very limited program due to his small yard, so I am left to see what I can pull out of them and I have the space to take these types of 'dirt roads' in my program. I don't expect each mating to give an introduction - only to advance the projects I am pursuing. They are a very interesting group of seedlings, and very consistent with little variation amongst them. I have culled some over the years, but the handful that are left are just exceptional plants. However, they do not merit introduction. Let's consider why. 

They are average, first and foremost. There are already introduced many little melon diploids that don't rebloom and have green scapes. That is the primary reason: they lack distinction - and what distinction they do have is in their exceptionally beautiful foliage - something not all that important, in and of itself without a distinct flower, to many daylily people (though it is very important to me). So there is just nothing there that hasn't already been done over and over, but what is important about these plants is the genes they carry.

Now, let's consider some of their good traits. First, they are all hard dormants and then, of course, the foliage - just look at it! They are so deep green and are not effected by adverse conditions. Look in the pictures how much darker in tone they are than some of the other seedlings and cultivars growing around them. The bed they are growing in is one of my worst test beds. It has awful sand/clay mix refill soil. The bed can get bone dry being on a slope, and yet, even with the mini-drought we had through May, the foliage is still gorgeous, no browning at all and no bud drop, unlike a good few other things here. Then they have a high scape to fan ratio and the scapes have an average of three branches plus a terminal Y on almost all of them and around 15 to 20 buds per scape and are about 24" to 28" tall. Most of them have the flowers carried just above the foliage and make a nice look in the landscape, especially as edgers. You will note in the pictures that there is one seedling that shows partial dark scapes, but it is the least attractive plant with the slowest increase and the least attractive habit and flowers (though that is in comparison to its sibling - it isn't really all that bad...compared to many other things). One seedling is closer to white than the rest and is a lovely flower. One seedling put up a few rebloom scapes in a really wet year, but has never done so before or since, so we know all the component genetics are there (we know that too from the parentage and knowing the behavior of the genes involved). It is now just a matter of combining those genes into homozygous expression...and then we might have an intro. Finally, these seedlings that are left have all been rust resistant for the four years I have had them, and I have tried to infect them every year (some of their siblings weren't rust resistant, and they are no longer with us...unless you consider the compost they made...)

So how am I using these seedlings? I am crossing them amongst themselves to bring the recessives back to homozygosity. That is, of course, an obvious thing to do. If I cross them back to pod parent E&O, I might get a higher number of rebloomers, but I won't get the dark scapes. If I breed them back to pollen parent Army of Darkness, I might get a higher number of dark scapes, but I won't get rebloom, so the only route to get both, fully expressed, in some small number of the seedlings is to cross the F1 with each other to make a true F2. As I am not necessarily looking for near white flowers, I am not calculating for that, and any shade of melon will work, and since these F1 seedling are all variations of melon, I can expect 100% melon offspring, in various shades, and any shade will be fine with me if it reblooms and has dark scapes.

Because these seedlings are all full siblings and all show the nice foliage that can handle dry conditions and poor soil and still look good, I don't expect a lot of segregation on this level. Both parents also have good foliage, so that suggests there will be little variation in that trait. All have good branching (for this type of thing - most of the rebloom dips have poor branching - usually just a terminal Y or W), so again, that should be fairly consistent in the F2 offspring. But, you may ask,"What if it isn't, or what if you get some poor foliage, etc?" Good questions!

In this instance, if such occurs with any desirable trait, then I can backcross any reblooming, dark scaped F2 seedling to any of these F1 seedlings. They have no obvious, serious flaws, so a backcross can be considered, and all the genes I want to see expressed are present in all these F1 seedlings. Further, both the F1 x F1 sib matings and the F2 x F1 backcross can reveal any hidden deleterious or undesirable genes that may be lurking in the line unseen, and that too is valuable information from a breeding standpoint. As well, a backcross may even concentrate the desired traits, both the rebloom and dark scape traits as well as the plant habit traits that are also very impressive.

This type of bridge plant project is very different from the one I discussed last night. This is not a salvage project, where you wish to incorporate good traits while removing bad traits. That is a much more delicate procedure. This is a simple breeding project to retrieve recessive genes. This is a very common reason to use bridge plants, perhaps the most common, and is how desirable recessive traits can be combined into one individual where they do not already occur together. In this instance, backcrosses are not just for test matings to determine when the undesirable traits are gone from any given individual in the line, but can also be used to concentrate desirable genes within the line.

Bridge Plants Part 3: Working from Tender to Hardy
In this post I want to talk a bit about working from a tender evergreen that is not hardy in my garden toward a hard dormant that is northern hardy.

The flower in the attached pics is Kanai Sensei x Blue Oasis. The cross was made in summer 2011. It is the only seedling I kept from the cross out of about 120 seedlings. It had the least offensive foliage in spring to my eye and taste and has been rust resistant over several years. Just by chance, it also carries the blue (bluish/lavender) eye trait of its pollen parent and shows some level of pattern, which was really lucky! It is what would be called a semi-evergreen and has fast increase and some rebloom.

However, it is not, to me, an introduction. It regularly curls at least one petal inward as you can see in the picture below, and sometimes all three. The foliage is not attractive to me and even though it was the only seedling selected, I am still not happy with how it looks in the spring - it was just better than the rest, but not really good.

I bought Blue Oasis in fall of 2010. The winter of 2010/2011 was mild and so it did well and bloomed nicely in 2011 and I used it that year as a pollen parent and pod parent (I also have one plant left from it as a pod parent - a story for another day...) The winter of 2011/2012 was also mild and the plant survived well, but we had late freezes and the foliage looked horrendous, bloom was suppressed and the clumps reduced fan count (there were two clumps of it). One clump looked much worse than the other and was removed, while the second clump was moved to my mother's garden. My mom lives in the old farm house my grandparents lived in and the soil around it is just amazing - black and rich. I consider her garden the remedial garden - if it won't grow anywhere else, it will usually grow in her garden... The winter of 2012/2013 was mild, but we had a series of late freezes and again, Blue Oasis dwindled further. Then came the winters of 2013/14 and this last winter, 2014/15 and Blue Oasis dwindled significantly each year. It is almost completely gone now, just two half-pencil sized fans from an original clump of twelve fans. One more hard winter and it will be gone, I fear. This is the common pattern for tender plants here in my area. They don't tend to just die here, though I have had it happen, but what is more common is that they dwindle away over a few years. If it couldn't survive in my mom's garden, it just doesn't survive well here. Mike Huben also told me that not only usually survive in his Boston area garden, but it won't bloom there either, even after milder winters. So I am very grateful that I got something out of it to move forward with, because it is really beautiful.

I have known since I started hybridizing that I wanted one aspect of my program to be moving fancy but tender faces toward dormant foliage, so I had chosen Kanai Sensei as a mate based on the foliage dormancy, the near white color and the strong increase and rebloom it shows. That was not an act of wisdom though, but a lucky break. I had no idea BO would dwindle as it has, or that KS would produce such good results in breeding. Sometimes you just get lucky by accident, and never underestimate the workings of lucky accidents in a program - they happen for all of us smile emoticon

So back to this seedling - There are some good traits here and the traits I don't like are really more about personal preference than an actual defect, so this is one of those projects where I am just not satisfied with this first generation result and want to keep working to get to a plant that is more what I want and what I want to introduce.

Many people are not bothered by the rolled inward or 'canoed' petals, but it irritates me, and that is just a personal preference. It is no reflection on anyone who likes that or grows or introduces plants with it, but I am obsessed with very open, very flat flowers - blame Richard Norris and his wonderfully flat Substantial Evidence family line! smile emoticon
Blue Oasis

The foliage, both the look of it and the habit, don't satisfy me and I have no idea how much further north beyond my area this individual might be hardy, and I want the bluish eye on hard dormants. Again, no reflection on anyone who likes other foliage types, but I am just in love with hard dormants. That is a personal thing, but with the tenderness of Blue Oasis, I would not feel comfortable saying this plant was a hardy semi-evergreen or evergreen. That again is just a personal inclination.

The plant does put up a rebloom scape or two most years, but it is not enough to call it a rebloomer and my goal is this look on real, consistent rebloomers like its pod parent. As well, there is poor branching and low bud count, as is common on many of those little rebloomers, which would be ok with me if it rebloomed heavily, but it doesn't. So all these things add up to something I want to continue on with, but that I would never see as an introduction in and of itself. It might make a great plant further south, but I don't live further south and I have to make my decisions based on what I see in my own garden. In my own garden, other than the flower, there is nothing to recommend this plant, other than the genetics it carries and the potential it offers for the future. It is the nature of breeding that sometimes we have to make bridge plants to get where we want to go. When material isn't available where all the traits are already combined, sometimes the only way to get there is to make the generations that you know won't give you want you really want, but will give you the pathway to it. That is one common area where bridge plants are a necessity, or a necessary evil, depending on how you look at it.

Kanai Sensei

So what am I doing with this plant? Well, I am actually using it in several ways. I have backcrossed it to Kanai Sensei and Early & Often, as both will allow eyes to express in their offspring and will help to concentrate the rebloom traits while keeping the nice, clean background cream color on the melon base. Then I am crossing it into seedlings from the Substantial Evidence family crossed to reblooming lines, both to flatten out the petals and to work to concentrate rebloom. Finally I am doing a few selfings of this seedling, just to see what happens and if I can pull out any dormants while also concentrating the eye trait and the rebloom trait. Odds are slimmer in the selfing of getting strong rebloom or dormancy, but it is possible. 

I suspect that what I will get from all the various breedings are more 'parts' to move forward with. I won't be upset in the least if I just get more bridge plants to continue weaving together into what I actually want. That is just breeding and when you are working to combine several traits that don't already exist in one plant, then it can take a few generations to put it all together into one plant, and that plant then be worthy of introduction. Of course, I might get something really nice out of one or all of those matings and maybe even have an intro, but I am never looking for that. I am just looking to extract traits I desire and maybe concentrate them - incremental steps.

If you look at the pictures below, they are mildly blurry because I am using my tablet right now, as the SD card went out in my good quality camera, but I plan to get one tomorrow, so I may add some more pictures of this seedling later on, but for now, these are good enough to see the flower and the eye. Note how the eye is patterned in bands of color as well as a bluish/purple/lavender. It is especially apparent on the sepals. The picture where I am holding back the petal for a better view especially show the layers of tones.

All in all, this is a nice seedling and not a bad plant. It might be a much better plant in a warmer area. It is close to what I want, but just not quite there yet, so onward we go, ever searching for the ideal. This is not a bridge plant because is has a defect, nor is it a bridge plant because it is boring (though there are many semi-evergreen small flowered cultivars with a similar eye by now, so it isn't terribly distinct), but because it doesn't meet my goal. I am satisfied that through it I can pull the beauty of the tender Blue Oasis into the hardy reblooming lines that Mike Huben has developed. Time will tell if I am successful wit the effort, but even if I am not, I have already learned a great deal through working with this plant and can apply that to other similar projects in the future.

Bridge Plants - Part  - Addendum 

In a recent post on bridge plants I discussed the plant pictured below (Ashwood Wray of Sunshine x Pigment of Imagination). I noted that it had pollen infertility, but several other very nice traits. However, in evaluating for thrip resistance this year, I found that this plant had very high thrip susceptibility (as does its pod parent), so I disposed of it. One bad trait is one thing, but two is too many for me in this instance. This plant was what I call a 'salvage project', but such projects have to be approached sensibly and with a realistic understanding of what a salvage project really entails. They are a lot of work! There is no guarantee that you will ever get anything out of a salvage project. You are always taking a gamble, and the more traits you are trying to eliminate/add, the greater the number of seedlings you have to raise and flower out and expose to various conditions in order to find the breakthrough plant you are after. So I am very cautious about salvage projects and I do not approach them lightly. I have a few seedlings from this plant already, and they will be watched very closely over the next couple of years, but I don't have high hopes for them, though you never really know. 

Since I had mentioned salvage projects and this plant in particular, I wanted to update everyone one its status and illustrate that selection sometimes means giving up on something you had hopes for. It was very difficult for me to make this decision, something I actually agonized over for several days, but in the end there was really only one decision I could live with. There is a bright side to everything if you choose to look for it. There are no tragedies or traumas, just opportunities if you choose to use them. Those opportunities can greatly improve your lines too, if you let them.

Friday, July 3, 2015

An Odd Season and Observations So Far...

An Odd Season and Observations So Far...

The 2015 flower season has been strange so far. Some things are very early, others late. Bloom time just seems to be off for many cultivars (and seedlings) and the weather patterns have been odd again this year. After a hard, cold winter with heavy precipitation, we became very hot and dry, very quickly, in the beginning of May. This brought on both early scaping and heavy thrip populations.

Many of the early scapes had been slightly frost damaged and were much shorter than usual, often way down in the foliage. The thrips destroyed many buds and even whole scapes on a lot of the early things, but in amongst those things were individual seedlings and cultivars that showed seeming resistance to the thrips, having very little damage or bud drop, often even surrounded by plants showing high susceptibility. Where there were many individual clumps of these seemingly resistant plants, there was consistent apparent resistance on all the clumps of those types, while where there were many individual clumps of the susceptible things, they all showed high susceptibility. Further, certain family lines showed consistently higher resistance or higher susceptibility. This likely indicates an actual mechanism that suppresses the thrip activity and may well be genetic. 

As those who know me or regularly read this blog will know, I cull for disease resistance, so the thrip invasion was a blessing, allowing me to identify those that showed seemingly resistant responses to the thrips. I culled as many seedlings in the early season as I typically do in the fall for rust selection. While it was hard work and a bit nerve-wracking at times (it takes a lot of intense observation and thought), it is work I am very glad to have been able to accomplish.

A highly susceptible daylily seedling showing severe thrip damage as blighted and browned scapes with total bud drop.

Of course, in our display gardens (which I tend not to breed form much), spraying for the thrips is the only real response (unless we want to dig out all the highly susceptible and replace them with resistant individuals - a thought for the future...), but in my hybridizing garden and seedling beds, I just can't bring myself to hide the problem and then pretend it doesn't exist, with a shrug of the shoulders and the comment, "Well, everyone should be spraying anyhow". I have way too many years of working with immuno-genetics to possibly think that way. I have also noticed that some very vigorous, hardy and wonderful cultivars coming from programs with a spray program have intense susceptibility to thrips (and usually rust too). This is because the seedlings are not being screened for resistance to thrips (or rust). It is a double edged sword. I understand why people don't want to go through the hassle of culling for resistance, but I also understand the importance of this type of selection. While we daylily fanatics might be happily willing to spray for problems, the average gardener will usually not. Of course, maybe we daylily people have become so insular that we don't care about average gardeners anymore? I do, but maybe some don't? I hope we all do.

This is a thrip resistant daylily that is literally swarming with juvenile thrips (increase the size of the picture to see them). They are like tiny grains of rice and are all over the petals, yet this flower shows little to no spotting from the thrips and little to no bud drop.

In addition to the selection for thrip resistance, the dry spell also gave the opportunity to select for drought tolerance. It is interesting to me how much variation there is in this factor in the Hemerocallis. Some cultivars remain lovely and don't even brown much, while others become heavily stressed and have the foliage brown and flop over into messy whorls of wilted foliage. I always observe and do some culling amongst the seedlings for drought tolerance in dry spells.

By the end of May we started getting rain again. By this time, many scapes were appearing and oddly, many cultivars and seedlings from across the whole season-range were scaping and preparing to bloom, often very much earlier than usual, though a few individuals have done just the opposite. I think the extreme heat and dry of May triggered a lot of mid/late-late plants into thinking it was July/August in May. The advantage to this was that I was able to identify some of those for thrip resistance or susceptibility during that hot, dry period.

So for most of the month of June and now going into July, we have been cooler and wet. In fact, we are now getting almost too much rain. So this will allow a whole range of different problems and thus selection to occur. We have seen this pattern for a couple of years now - hot, dry spring and cooler, wet summer. It makes for an odd mix of conditions and selection possibilities. I always say that there are no bad conditions, just opportunities for selection.

This has also been an interesting year for me in that it is now my fifth year of actively breeding daylilies, though I have grown daylilies for forty years as of this year. So I am now seeing some mature seedlings in fully clump-strength that have been selected for that whole time period for a wide range of characteristics, and finally just this year they are getting culled for flower traits. 

My program is based around developing my own lines and hopefully some unique styles that are not being focused on much by other breeders. To do that, much of my early breeding revolved around crossing widely different types to bring different traits together, and so that produced many plants that are simply bridge plants. I am very excited about much of what I am seeing from these seedlings, now in their fifth year. Some are actually further along in the phenotypes I wanted to go toward than I expected.

One thing that is different about this year is that I find myself using my own seedlings much more heavily in hybridizing than in any year in the past. It struck me just a couple of days ago that when I am pollinating within certain projects, my first thought now is to go to certain seedlings for pollen over any cultivar. That is an exciting stage, and from many years of breeding animals and plants, I recognize this stage. It means I am moving into my own lines. I am not away from other folks intros yet, and I probably never will be completely (I use what's good, no matter where it came from or whether I or someone else bred it), but it is an exciting  point to realize that this is unfolding before my eyes and I am seeing some very interesting things just beginning to emerge. It has allowed me to refocus my intent and to feel very encouraged and excited about what I am doing.

Breeding is both an art and a science. There are many, many methods that will all work. I don't use just one method or another. As both an experienced breeder and a former poultry researcher and science-obsessed-geek, I know many methods and techniques for breeding and I use what I think may work best in any instance. I don't put myself in a little box and refuse to look outside. I want to use what works, not give allegiance to one or the other method. Methods are merely vehicles, they are not the actual journey...

I often see people, well-meaning undoubtedly, trying to force everyone into 'their method'. People will probably think I have a 'method' also, but I really don't. I just use what works, and I know what works from many years of reading, research, experimentation and experience, but each cross is always an experiment, except perhaps where you have worked with both parents in the cross extensively and so know them very well, and even then you may well get surprises. The notion that you must only use new cultivars in breeding is perhaps more about hybridizers selling new cultivars than it is about any genetic reality. I do think some hybridizers seriously believe that only the newest cultivars are worth breeding for, and that may in fact be true if the only interest is in making the greatest advancements on already exaggerated traits in the shortest possible period of time.

Purchase price of a daylily is no reflection of its breeding ability or its genetic content. As well, low priced and older plants are not by default 'bad' or 'useless'. That is ideology, not genetic fact. Many older plants have much to offer. Many were not used well and we have much more advanced genetic material to combine with these older things now than the breeders then had available, so looking at what was produced from a given plant fifty years ago does not tell you what it can produce now when combined with a modern cultivar. 

Good examples are rust resistance or branching. Many older cultivars offer these traits and can be used to bring these traits into more modern lines. You might only get a bridge plant in the F1 (first generation), but the use of bridge plants is a well-known and well-used technique in all plant breeding. We owe many of our finest cultivars to bridge plants. For instance, Substantial Evidence and its descendants and relatives would not exist without bridge plants, because as Richard Norris points out on his website, the F1 (and F2) from Lights of Detroit x When I Dream (both "old" cultivars at the time Richard used them...) were not much to look at, but he persevered with them and look what we have now because of it! There is not just one method. Don't believe anyone who tells you there is and question what they have to say if they insist there is only one way to go...That's not to say that their methods are 'wrong', just that there may be many other ways that can also work. 

In closing, I want to encourage everyone to follow their bliss, pursue their visions and experiment. There are no mistakes or failures. There are only opportunities to learn and grow. Do not always throw away F1 plants if you don't get an intro (always remember, there are these things called 'recessive genes'!!). Don't overburden yourself with too many bridge plants, but having a few around is not a bad idea and can give you some amazing breaks in the long run. Think about plant traits and disease, pest and environmental resistance as much as you think about the face and you will produce more well-rounded offspring. Don't let anyone tell you something is not possible or that you are wrong for pursuing what you want in your own program. The only arbiter is the genes. They may tell you something is not possible, but that is something you can only discover from making the attempt.

As usual, there will be fewer blog posts here throughout the summer and into early fall, but on rainy days like today, I will make every effort to get a post out every month or so. Enjoy your plants and pursue your dreams!