Monday, November 5, 2012

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.