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Jun 12, 2018 4:21 PM CST
Thread OP
Alabama
So, im finding confusing information regarding the species fulva, I have never personally seen a diploid var, fulva. but i have heard that they do exist..

Daylily
'Hankow'
is diploid and fertile? what is the origin of these diploid varietys and how many exist? I also found this article.




(Hemerocallis fulva ex Korea, Seoul National University, NA 54920) was collected by Darrel Apps and Barry Yinger on a 1984 collection trip to South Korea and is not a registered cultivar, as some fulva clones are such as 'Hankow' (Stout, 1939) .

In an article in The Daylily Journal Vol. 43, No. 1 - Spring, 1988., in an article titled Korean Daylily Species by Darrel Apps and Lynn Batdorf is found this description of this fulva clones, on page 18, "Adventuresome hybridizers might be interested in one other plant found in Korea. Time after time in cultivation we saw a plant much like H. fulva 'Europa'. However, we also kept seeing a similar fulvous daylily of a brighter color. At Seoul National University we found what appeared to be a fertile form of H. fulva (the cultivar 'Europa' does not set seeds). This plant is labeled NA 54920. Its flower scapes were 46" tall and the flowers up to 5" in diameter. The throat of the flower was a clear yellow. It had a deep red eyezone and petals and tepals were orange, but with fulvous red veins. Although no attempts have been made to pollinate the flowers, some have formed pods and seem fertile."

Also I just thought id share.
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Jun 12, 2018 6:01 PM CST
Name: Larry
Enterprise, Al. 36330 (Zone 8b)
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Jun 12, 2018 8:52 PM CST
Name: Deborah
midstate South Carolina (Zone 8a)
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Avatar for clockworkhawkins
Jun 12, 2018 9:03 PM CST
Thread OP
Alabama
thanks guys Smiling any information or links regarding unreduced gametes and their relation to H. minor. are appreciated. also anyone with experience with colchicine. id greatly appreciate your input and information , I've read misleading information and would like to clear it up
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Jun 13, 2018 6:19 AM CST
Name: Diana
Lincoln, NE (Zone 5b)
Daylilies Region: Nebraska Organic Gardener Dog Lover Bookworm
This sounds like a post for Maurice @admmad
And Welcome! : )
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Jun 13, 2018 7:38 AM CST
Name: Maurice
Grey Highlands, Ontario (Zone 5a)
@clockworkhawkins

The natural species of Hemerocallis are diploids. They can still be found growing in various parts of Asia. If a single plant of one of those natural species is dug from the wild and grown in a garden then it can become a cultivar and given a cultivar name.

'Hankow' should have originally been a wild plant of the species Hemerocallis fulva that was collected and grown in gardens. If it was simply one plant growing in a meadow or field where many other H. fulva plants were growing, then it probably would be a normal diploid. On the other hand, triploids can rarely occur naturally and since they are different from normal plants (sterile, do not produce seeds normally, etc.) they sometimes are specifically chosen (the plant that is different from others in a field - it sticks out) and brought into gardens.

Although unreduced gametes are discussed in daylilies, the objective evidence suggests that they are extremely rare. In species that produce unreduced gametes at useable rates one tends to find that the natural species are not just diploids but also tetraploids, or above. There are some natural triploid daylilies but no natural tetraploid daylilies suggesting that unreduced gametes in daylilies are very rare. I do not know of any evidence that suggests H. minor produces unreduced gametes. Arisumi, a geneticist, conducted research looking for unreduced gametes in cultivated daylilies - he did not find any and made a large number of crosses in that search.

What is the misleading information about colchicine?
Avatar for clockworkhawkins
Jun 13, 2018 2:50 PM CST
Thread OP
Alabama
I spent 6 hours last night, reading through your posts and others articles regarding nonreducing gametes. thanks so much for the reply.

the cultivar "implausibility" is extremely interesting , but I am confused . how can a triploid even create a gamete how many pairs does it create?
Nick Chase felt it was a 2N gamete from 'Europa' that combined with an unreduced 2N gamete from Ed Murray that produced a 4N tetraploid offspring. but what happens at the other pole?

3/2=n/a
just focusing on fulva the egg is some how a 2n? do triploid always give off 2n? or can it be haploid just one set?

Are there anyother daylilies with claimed unreduced gamete production besides ed murray?

did Arisumi test ed murray?

another note.

from the article
RESEARCH ARTICLE
Ploidy variation and karyotype analysis in
Hemerocallis
spp. (Xanthorrhoeaceae)
and implications on daylily breeding

Most Chinese cultivars were diploid; only one was tetraploid. Among the 29 wild genotypes
collected from the Taihang Mountain range, 13 (45%) were triploid and 16 (55%) were diploid; no tetraploids were identified.

given that wild triploids do occur in nature. I would assume give an isolated population. of both dips and trips . Unreduced gametes would have to exist. how else would we have trips naturally?

I should have used the the word conflicting not misleading in regards to colchicine.
Ive read multiple clinical trials that suggest that it is not carcinogenic however, its has been suggested to me many times that in fact it is.

also. Im trying to track down the first hose-in-hose daylily . but i could only find double golden eagle from 48 I believe?
Last edited by clockworkhawkins Jun 13, 2018 5:45 PM Icon for preview
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Jun 13, 2018 3:33 PM CST
Name: Sue
Ontario, Canada (Zone 4b)
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Hemerocallis fulva 'Flore Pleno' could be considered the first known hose in hose I would think. Another triploid.

Avatar for clockworkhawkins
Jun 13, 2018 5:49 PM CST
Thread OP
Alabama
do we know if its caused from a duplication of petal alleles in the dna? i was thinking there was a possibility of interference during somatic doubling that caused the section to show up twice ?
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Jun 13, 2018 6:03 PM CST
Name: Maurice
Grey Highlands, Ontario (Zone 5a)
@clockworkhawkins
"how can a triploid even create a gamete how many pairs does it create?"
In a diploid the chromosome pairs can align without problems as they each have only one partner.
In a triploid the chromosomes can align as triplets or pairs with singles. Each of the 11 sets of chromosomes in a diploid can align independently from all others. The end result is that the gametes have a wide range of chromosome numbers in a triploid. However not all chromosome counts in a gamete are viable. So the range of chromosome numbers in viable gametes produced by a triploid are much narrower. The normal chromosome count for a diploid gamete (eg pollen) is 11, the normal count for a tetraploid is 22 but a triploid might produce gametes with 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28, etc. chromosome counts. Most gametes produced by a triploid are not viable. Most embryos produced by crosses involving triploids are not viable or the endosperm (nutrient tissue in the seed) is not viable or both. So most triploids in most species are considered to be sterile. In some cases, with enough crosses one may get lucky and have a viable seed(s). Arisumi tended to find that triploids were more fertile with tetraploids than they were with diploids.

No one has checked the chromosomes of 'Implausibility'. There is at least one known triploid (chromosomes counted) that is quite fertile with tetraploids. It is plausible that 'Implausibility' is simply another triploid (or other type of aneuploidy). There is more than one plausible explanation for the fertility of Implausibility with tetraploids. Aneuploids have unbalanced chromosome counts. So a diploid daylily should have 22 chromosomes (11 pairs - two of each set). An aneuploid may have 20 or 21 or 23 or 24, etc. chromosomes (only one chromosome of some sets or three chromosomes of some sets, etc.).

A triploid would (normally) extremely rarely produce an "unreduced gamete" (unreduced does not really apply to the triploid gametes - 2n gamete is the better term) of 22 balanced chromosomes. Whatever frequency it produced such gametes should be equal to the frequency at which it produced balanced gametes of 11 chromosomes and therefore be capable of producing diploid seedlings when crossed with a diploid. That probability might be estimated from the number of crosses Stout made with 'Europa' and the number of viable seedlings he produced. However that assumes that each one of those seedlings had 22 balanced chromosomes and that is unlikely.

Arisumi did not check 'Ed Murray'.

fulva 'Korean' has been claimed to produce 2n gametes. There are difficulties with that idea.

"Ploidy variation and karyotype analysis in Hemerocallis spp. (Xanthorrhoeaceae) and implications on daylily breeding"
This research has unknown problems that resulted in the researchers "finding" that a number of cultivars have a different ploidy from that registered or from that determined by other researchers. I consider their findings unreliable.

However, it is known that triploid daylilies exist naturally. Triploid plants can be created naturally by somatic chromosome doubling. It is not known (as far as I know but I have not done a literature search) what proportion of natural triploids were created by somatic doubling (or any other method) in comparison to unreduced (2n) gametes.

Colchicine changes the chromosome number. Almost by definition that may result in cancer if it sometimes produces some cells with unbalanced chromosome counts. It also causes other changes and probably also causes mutations of other sorts. However, it is apparently still used medicinally.
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Jun 13, 2018 6:08 PM CST
Name: Maurice
Grey Highlands, Ontario (Zone 5a)
In most cases studied flower doubling is caused by mutations in specific (sometimes known) genes. Sometimes the mutations are "dominant"; sometimes they are "recessive" but they probably typically show variable expressivity (they do not produce the same characteristic each time) and they may have incomplete penetrance (individuals may have the mutation and should show double flowers but do not).
Avatar for clockworkhawkins
Jun 13, 2018 8:41 PM CST
Thread OP
Alabama
as in being phenotypical? I found their research regarding the idea that triploids may be the perfect or more perfect chromosome count very illogical. they based their findings off the idea that these triploids were larger. but they weren't testing a trip and a tet of the same genetics. to me that's not a control at all. so lets leave the realm of triploids and say I want to isolate diploids that are believed to create 2n gamete if I then pollinated them to a tetraploid and line bred from there could I end up with a tet many years down the line that produced 4n gamete?

I've heard somatic double can be induced through stress during the development of buds such as temperatures above 110f?
Last edited by clockworkhawkins Jun 13, 2018 8:44 PM Icon for preview
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Jun 13, 2018 10:50 PM CST
Name: Diana
Lincoln, NE (Zone 5b)
Daylilies Region: Nebraska Organic Gardener Dog Lover Bookworm
... Gentlemen, the pair of you read like one of the masters classes I'm taking this summer. Different subject matter, but brain-busting all the same! I hope you both carry on- this is quite the education! I tip my hat to you. Thank You!
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Jun 14, 2018 6:29 AM CST
Name: Maurice
Grey Highlands, Ontario (Zone 5a)
@clockworkhawkins
clockworkhawkins said: lets leave the realm of triploids and say I want to isolate diploids that are believed to create 2n gamete if I then pollinated them to a tetraploid and line bred from there could I end up with a tet many years down the line that produced 4n gamete?

Assuming that the ability to produce some 2n gametes is genetic then yes. In other plant species specific mutations have been found that cause errors that result in 2n gametes.

I've heard somatic double can be induced through stress during the development of buds such as temperatures above 110f?

Both high temperatures and low temperatures are known to cause 2n gametes. That would occur during the development of flower buds. Temperatures of 110F (42-43C) during the development of seed have caused somatic doubling. The high temperature was applied as a temperature shock lasting 20-30 minutes at the time of the first division of the zygote. Treating apical or axillary meristems during the development of vegetative buds with very high or low temperature shocks may produce chromosome doubling. Twin seedlings sometimes have polyploid (more sets than normal) chromosome counts.
Avatar for clockworkhawkins
Jun 14, 2018 7:33 PM CST
Thread OP
Alabama
admmad said:@clockworkhawkins

Assuming that the ability to produce some 2n gametes is genetic then yes. In other plant species specific mutations have been found that cause errors that result in 2n gametes.


Both high temperatures and low temperatures are known to cause 2n gametes. That would occur during the development of flower buds. Temperatures of 110F (42-43C) during the development of seed have caused somatic doubling. The high temperature was applied as a temperature shock lasting 20-30 minutes at the time of the first division of the zygote. Treating apical or axillary meristems during the development of vegetative buds with very high or low temperature shocks may produce chromosome doubling. Twin seedlings sometimes have polyploid (more sets than normal) chromosome counts.



I was under the impression that the 2n gamete developed via pollen during the first stages of bud development? Correct me if im wrong but if the pollen develops as normal it produces a haploid. but are you saying you take the haploid pollinate the flower then crank the heat to 110 for the first half hour? Smiling I have my son for the next few days so I may be slow to reply. thanks very much for your information its been very educational. I have a few more questions ill be in touch when able
Avatar for clockworkhawkins
Jun 14, 2018 10:12 PM CST
Thread OP
Alabama
admmad said:@clockworkhawkins

Assuming that the ability to produce some 2n gametes is genetic then yes. In other plant species specific mutations have been found that cause errors that result in 2n gametes.


Both high temperatures and low temperatures are known to cause 2n gametes. That would occur during the development of flower buds. Temperatures of 110F (42-43C) during the development of seed have caused somatic doubling. The high temperature was applied as a temperature shock lasting 20-30 minutes at the time of the first division of the zygote. Treating apical or axillary meristems during the development of vegetative buds with very high or low temperature shocks may produce chromosome doubling. Twin seedlings sometimes have polyploid (more sets than normal) chromosome counts.



I was under the impression that the 2n gamete developed via pollen during the first stages of bud development? Correct me if im wrong but if the pollen develops as normal it produces a haploid. but are you saying you take the haploid pollinate the flower then crank the heat to 110f for the first half hour?

have you heard the terms triploid block and triploid bridge? I've read that its been suggested that the triploid is the next stage in evolution. that its essentially waiting for the correct gamete formations. at first i thought it was plausible but now I disagree.

I assume a nonreducing gamete flower needs to be met with other 2n gametes to preform the switch to tetraploid. however if the triploids are spawned from 2n x 1n = 33 pair then they would also be carrying atleast the genetic probability for 2n given to it by 1 of its parents.

my next assumption delves into the realm of science fiction. given a perfect series of line breeding where a large gamete formation was not just plausible but the norm rather than the rarity could we assume that perhaps a dodecaploid giving off 12n could actually hybridize higher up its classification. a theoretical example being.

two plants being 12n gamete in the family Asphodelaceae. obviously you have genetic misconfiguration but given a larger gene pool would there not be a higher likely-hood of success?
Last edited by clockworkhawkins Jun 14, 2018 10:27 PM Icon for preview
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Jun 15, 2018 6:37 AM CST
Name: Maurice
Grey Highlands, Ontario (Zone 5a)
clockworkhawkins said:
I was under the impression that the 2n gamete developed via pollen during the first stages of bud development? Correct me if im wrong but if the pollen develops as normal it produces a haploid. but are you saying you take the haploid pollinate the flower then crank the heat to 110 for the first half hour?

Natural polyploids may be produced by different methods. They may be produced by 2n gametes or by somatic chromosome doubling.
2n gametes may be male or female and may be produced by specific mutations or by environmental effects (temperature shocks). They may be produced at two different stages of meiosis in flower buds.
Somatic chromosome doubling may be produced by environmental effects (temperature shocks) at early stages of shoot apical meristem or axillary meristem development in vegetative buds.

If you pollinate a diploid flower with normal pollen and know the time at which the embryo undergoes its first division and apply a temperature shock then you sometimes may cause somatic chromosome doubling and that seed may develop into a tetraploid. The timing would not be in the first half hour; it is not known for daylilies.
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Jun 15, 2018 7:08 AM CST
Name: Maurice
Grey Highlands, Ontario (Zone 5a)
clockworkhawkins said:
have you heard the terms triploid block and triploid bridge? I've read that its been suggested that the triploid is the next stage in evolution. that its essentially waiting for the correct gamete formations. at first i thought it was plausible but now I disagree.

I assume a nonreducing gamete flower needs to be met with other 2n gametes to preform the switch to tetraploid. however if the triploids are spawned from 2n x 1n = 33 pair then they would also be carrying atleast the genetic probability for 2n given to it by 1 of its parents.

my next assumption delves into the realm of science fiction. given a perfect series of line breeding where a large gamete formation was not just plausible but the norm rather than the rarity could we assume that perhaps a dodecaploid giving off 12n could actually hybridize higher up its classification. a theoretical example being.

two plants being 12n gamete in the family Asphodelaceae. obviously you have genetic misconfiguration but given a larger gene pool would there not be a higher likely-hood of success?


Triploid block is often mentioned as being present in daylilies. It is not present. When a triploid block is present in a plant species crosses between diploid and tetraploids do not produce triploids but rarely produce tetraploids. So in a species with a triploid block all (or more reasonably nearly all) seedlings produced from crosses of diploids with tetraploids would be tetraploids. In daylilies Arisumi showed that crosses between diploid and tetraploid daylilies rarely produce seedlings but when they do they are triploids not tetraploids.

Triploid bridges are considered as one way for natural tetraploids to be produced. The standard scientific consensus was that triploids were sterile. However, although triploids are more or less sterile they are not necessarily 100.0% sterile all the time or in all species. Therefore it is now considered that natural triploids may be one possible mechanism to produce natural tetraploids.

Tetraploids do not typically have normal fertility. Newly created higher polyploids presumably would have even lower fertility. Arisumi created one hexaploid . It was completely sterile in crosses with diploids, triploids and tetraploids and could not be crossed with itself.
Avatar for clockworkhawkins
Jun 16, 2018 11:50 PM CST
Thread OP
Alabama
thank so much for this information. I've learned so much
my final two questions.

are there pictures of this hexaploid?
what is the reason for loss of fertility?
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Jun 17, 2018 7:17 AM CST
Name: Maurice
Grey Highlands, Ontario (Zone 5a)
clockworkhawkins said:thank so much for this information. I've learned so much
my final two questions.

are there pictures of this hexaploid?
what is the reason for loss of fertility?


You are welcome.

There is a black & white photo of the flower in Arisumi's article. The article is CYTOLOGICAL, HISTOLOGICAL, AND MORPHOLOGICAL EVIDENCE FOR A COLCHICINE-INDUCED HEXAPLOID HEMEROCALLIS By Toru Arisumi and L. C. Frazier and is in the Daylily Journal archives. I do not have information on which issue.

In diploids each chromosome of the chromosome pairs has only one choice for pairing at meiosis. So they pair symmetrically at meiosis and there are 11 sets in daylilies. In tetraploids each chromosome has a number of possibilities when it comes to pairing/aligning. One set of four can align as a quadruplet or it can align as a triplet with one "unpaired" or it can align as two pairs. When the chromosomes separate, singles can be lost completely and quadruplets may separate as three and one rather than two and two. Each set of four does this independently of how the other sets align. The end result is that in diploids basically 100% of the gametes have a balanced chromosome set of one chromosome for each of the 11 sets - they have 11 chromosomes. In tetraploids the gametes do not necessarily have 22 chromosomes and even when the number is 22 it may not be two of each of the 11 sets. Unbalanced gametes are not necessarily viable and the seeds created by viable unbalanced gametes are not necessarily viable. The situation in triploids is much worse in terms of balanced chromosome sets in gametes. In hexaploids the alignments would be more variable and the viable gametes fewer.

Some research evidence also suggests that there are other factors that affect the fertility of tetraploids and higher polyploids. I don't know if there is any scientific consensus on what those other factors are and they may vary in different species.

It is known that one can modify the fertility of new tetraploids by selection. If one only breeds from those tetraploids that produce the most seeds then one can fairly quickly increase the fertility of the tetraploid population. Unfortunately that is not happening in the daylily tetraploid population and even if it was, since there are newly converted tetraploids being continuously created, it might not be effective.

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