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.
