Elizabeth,
Of all my hellebores, 'Mrs. Betty Ranicar', a fully double pure white without any spots , produces the best foliage. The flowers of other plants here range from almost-black to yellow, mauve, spotted, rose and white. By far, 'Mrs. Betty Ranicar', an orientalis type, exceeds all in both the amount of flowers produced and the clean shiny foliage.
From seed it comes about 90% true.

Hello Elizabeth and Carol ,

Elizabeth , the description of your test is almost professional , I do like it very much . So I hope you accept now a critical discussion .
The pictures that you refer to could be found in my Rice and Strangman book , a Dutch version of 2000 , on the same pages as you indicated .
( the pagination has not been translated ;-)

When I understand well your genetic formula , you are supposing that your Old Spot has the following genetic content ( AaBbCc ) and by selfing , the offspring gives all kinds of intensity of spots - and 1/64 unspotted .
Let us accept the 2/28 pure whites of your test as nature is not perfect , we also need more crosses to be able to really come to conclusions .
The other supposition you make is that the genes are (co) - dominant .

A correspondent from Holland reported me about his test : Spotted x Unspotted gave 50% spotted seedlings . It would be interesting to test your formula on this case but I did not because there is some other information we have to deal with .

Breeder over here told me that for Unspotted crosses they always become some spotted seedlings , up to twenty percent . ( twenty must be a max )
With the test I did , I became 15% .
If the genes are dominant , we can never get spots from an Unspotted cross , I think .
One thing I admit with you is that there must be some "distribution" at work , and as an unspotted cross mostly give some spotted seedlings ( the reverse is also true as you could see in your test ) there must be a point where spots pop up or disappear .
I was thinking at plastids , but a polygenic approach is also possible .
In both cases it is as if every spotting gen (or plastid ) that is present adds some effect , up to a point where we can see the spots .

Personaly I think that we must go for a combination of both plastids ( yellow ) and genes ( spots , black ) .

Carol , I found a very nice picture of H. Mrs. Betty Ranicar on the Online seed and plant catalogues of Thompson and Morgan (uk) .
This is a double white hybridus , so if you tell us that you got a 90% true breeding what are the others doing ? ( color , spots , anemonoid or else ) .

As I do not have doubles in my garden , I have to ask for information where I can .The genetic behavior of doubling might be very important to understand better what is going on with colors , there is probably no influence of the plastids in this case .
Up to now it is not possible to come to conclusion or even an attempt to it .

johan

PS for the moment I can't reach my homepage ,a hostingproblem , so I'm not able to put the new 2004 pic's and information . It is not a commercial site you know .

Here is a link that might be useful: fynwerk

Carol,

Wow! Mrs. Betty Ranicar is lovely. Like Johan, I am curious about what the 10% that don't come true look like? Please let us know.

Johan,

I'm grateful for your critical comments. I was in Amsterdam last September on a pilgrimage to the Hortus Botanicus. If I'd known you were there in Belgium, I would have tried to get together somewhere for tea and a good hellebore genetics discussion. Maybe someday!

Polygenic inheritance of spots does seem to make the most sense. As you say, "it is as if every spotting gene (or plastid) adds some effect."

I hope that there are only three gene pairs that control spotting, as traits in some organisms are controlled by as many as 100 gene pairs!

So, in my thought experiments on spotting, I am starting with three gene pairs, as I can handle the math for this.

I'm interested in the report from your correspondent in Holland that a cross between a spotted and an unspotted yielded 50% spotted seedlings. That would be the percentage we would predict if the spotted parent's genotype were Abc and the unspotted parent's genotype were abc, with capital letters representing spotting pigment and small letters representing the absence of pigment. Did your correspondent in Holland find that the spots were all of about the same density, or did they fall across a range of density?

It would be very interesting to know what crossing his or her unspotted offspring from this cross might yield. If the offspring from such a cross of unspotted parents included spotted forms, then my "abc + abc = unspotted" hypothesis does not work.

It doesn't seem to work for the breeders you mention who get around 20% spots from a cross of unspotted parents or your own experiment that yielded about 15% spotted forms.

So I am going to have to read up on plastids! Thank you for suggesting this route.

Two other factors that might be important to consider are: 1) whether we are all working with a common definition of unspotted, or if some of the unspotted forms actually have a few freckles here and there
2) if less spotting pigment, which comes from anthocyanins, renders seedlings and young plants more vulnerable to disease, so that there is a heavy attrition of unspotted forms before they reach flowering and can be counted. It's my observation that the whiter the flower, the more vulnerable it is to problems with fungus.

Can you think of a model that would account for the 15-20 or 25% of spotted offspring from a cross of unspotted parents? Three or more gene pairs with one or more genes dominant or recessive in relation to the others?

Thanks for helping me think this through.

Best,
Elizabeth

I asked Dan Hinkley and the others are single white. I have my first seedlings now ready to flower.

Dear Carol,

Thanks for the information! That's interesting. We'll have to add your information to the data bank and see if we can figure out the genetics of doubling.

It must be so exciting to see your the buds on your first seedlings. They will be gorgeous! Are they two or three years old?

Thanks again.

Best,
Elizabeth

Carol ,

Thanks for the information , I hope other people will join and give information on their seedlings . Up to now the information I could collect concerning doubling is a bit contradictory , especially the results with anemonoids . A German correspondent (breeder !) has the impression that the results are influenced by climatic conditions and a Belgian ( breeder ! ) wrote me that by crossing doubles he becomes all the way better results , the number ( % ) of double seedlings is growing .

Elisabeth,

The information of my correspondent of Holland was of great interest ;
He "estimates" 50% spotted seedlings of the cross ,
H.orientalis guttatus ( spotted mother ) x H.cyclophillus ,
but no information about the intensity .
He was doubting about the coloring by plastids , except for yellow .
There is no valuable , scientific , information on Hellebore colors .
The text he transmitted me confirmed that all hellebore have 32 chromosomes , all the wild forms and the studied gardenplants .

Now let us go for the color and spot problem ; It is correct to ask everybody to speak the same " color "-language .
My observation over several years is that flowers with some Rose ( by assumption : at least one gen for Rose ) darken and the spots are smaller when we have a cold spot over here ( Zone 8 ) .
Now the weather is very soft , so we see softer colors and bigger spots . Very important : My original Rose ( unspotted ) has some flowers with very small spots at the base of the sepals !

That is one of the observations that brought me to the plastid hypothesis as possible part of the coloring process .
The objection to that kind of observations is that this is probably not true for other climatic zones , or there might be some chemical influence of the soil ( ex Hortensia ) so it is not observable in other gardens . The point is ; can we make this observations more objective by measuring , calibrating and so on . In general an average gardener does not have the budget of a NASA moonwalker , but I'm sure it must be possible with some kitchenmaterial or by using light ( fluorescent ) or by putting a filter on the camera .
Observing the plants in a greenhouse , as I wrote almost a year ago , enables us to compare better year by year , but is probably not much better if we want to compare with other climatic zones except if we are not low budget gardeners .

About the protective effect of spots , anthocyanins .
Not all (sub)-species have spots or contain anthocyanins , so I think your observation seems not very correct for wild plants . Nevertheless I also observe some behavioral differences within my seedlings . On the garden web there were also some observations going the same direction .
The paler colors are more susceptible for stress ( they flower out of date ) and the seed is more endormant (for the seedlings that pop up a whole year after sewing the whitish are in higher number ) .
Perhaps I can mention the following observation here . Last year I had a big loss (80%) , only of my two year old seedlings in pots , after a very and exceptional cold spot . A honest breeder told me the same . What is the acceptable return when starting with 100 fresh seeds and cultivating unto flowering ? How to find out the relation between color and return ?

Concerning the polygenic formula you put forward , it is evident that it may not contain a dominant factor , to explain the spots .
We agree that we have to do with some " distribution " ( poly-something ) and that the spots pop up or disappear in some seedlings of an unspotted or spotted cross ( a yielding point ? ) .
By browsing some genetic books , I could not find a polygenic (diploid) example explaining such a result .
With my observations and also the observation of breeder that the motherplant is of great importance ( Attention they are breeding for colors , yellow and black !! ) I came to a model with plastids ( multicopies ) . The literature on this subject is moving and ... very difficult as the vocabulary is not stabilized !
Recently I found another possibility ( it is called paramutation ) found in Zea Mais . The anthocyanin is formed by genes and there are three possible alleles A , Asp and Am . You divine that "sp" means spot and "m" is marbled .
In the triploid context of the fruit , the alleles may be absent or present in different number . With six possibilities you can go from nothing ( white) to very dark red marbled and all possibilities between .
A lot to think about !
Perhaps I may give my impression on the last one . Just browse in the plastid literature and you will see that a lot is changing , not always accepted by the " classical " genetesists who only think " genes " .
I am always peplex that experienced gardeners over here can give me a lot of information without knowing anything of genes ( mostly I have to listen and to translate ) sometimes they just are speaking about cytoplasmic inheritance .

To set up a data bank is only valuable if there is information , but information on hellebore breeding seems very scarce . It may be successful if we could formulate questions so that average gardeners , as we all are in the garden , can also participate .

Your suggestion for tea somewhere between Brussels and Amsterdam is fully appreciated , but the distance between this cities is more than 200 km and we still calculate in cm over here .
I prefer to answer very politely ( translation ) : It is as if I became it !

Johan

Here is a link that might be useful: fynwerk

I have not found any significant disease susceptibility related to colour of flower or leaves or prescence/abscence of spotting. All species I've grown appear to be susceptible to botrytis in flowering stems and around carpels especially if the shed anthers don't blow off cleanly. Have been experimenting for some 10 years now.

The genetics issue is interesting as like everything else to do with helleborus it appears to very complex any highly variable - don't you just love the little possums! if it was all predictable would you still?. Just a gut reaction but I don't think its simple polygene control in spotting. I think there may be genes controlling position and size of pigment patches that we interpret visually as spots 0r blotches.
When does spotting become blotching? How can we explain star colouration at flower centre with no spots, with veining, with picottee? Howabout nectaries of different colours to the centre star same colour as centre star?

Doubles - well there's a question! 1 plant some flowers fully double - no nectaries, some part double part anemone some purely anemone. Early flowers more likely to be double than late. 1st year flowering one or to double flowers second more double flowers third year mailny doubles? all on the same plant!!!!!
Other plants double with nectaries others double fully without nectaries all double flowering from first year.
Oops! looks like environmental effects as well as pure genome or perhaps some genomes are more affected by environmental factors than others.
Sorry no answers just lots of questions and observations. unlike Mendel I don't have enough time to sit around doing stats analysis on thousands of crosses and then to fudge the results just ever so slightly to fit the hypothesis( sorry for the cynicism) - he did a good jod too.
I am genuinenely interested but have to make a living too sadly.

Johan,
Have just looked at your website - keep up the good work your english puts my french to shame!
Re: colour and changes due to environment have you researched at all the work done on the effects of environmental stressors on the metabolic pathways and enzymatic branching of same for the anthocyanins and flavenoids? I've found little specific to flowering plants and nothing on hellebores but the research on plant crops is interesting and provides hypotheses that support many of our observations on flower colour changes with season and climate.
regards Mike

Dear Mike:

Very glad to read your post. It's always wonderful to hear from someone with an experimental orientation. I'm typing this at the library, since my computer is in the shop (it spends more time there than at our home) and my time is running out, but I hope that we can keep sharing hypotheses.

Love all your questions re: picotee, spotting becoming blotching, etc.

I agree that it's unlikely simple polygenic control determines color. However, it's the most useful working hypothesis or paradigm that I can use for breeding right now. Having given up the luxurious simplicity of dominant/recessive patterns, I now expect that when I self an individual, I will get a range of offspring, some of which are likely to be twice as whatever as the parent, some of which are likely to be half as whatever as the parent, and many which are in between. I hope that makes sense. I'm getting better results now that I'm using this paradigm. But you and Johan are no doubt correct that there are many operative genetic variables.

Thank you for sharing the fact that you've never been able to find anything about hellebore genetics. Seems odd, doesn't it?

Re: anythocyanins and disease resistance. My records definitely indicate that my white hellebores are highly susceptible to fungus problems, while it is rare in my colored hellebores, and pretty rare in my more spotted ones.
This may, of course, be due to particular variables in my garden or in my care of them, or I may have been breeding a line of particularly susceptible white hellebores, in which their susceptibility has nothing to do with their whiteness.

Have you ever noticed that your white onions get fungus problems far more readily than your purples? I have.

I will try to find the citation on anthocyanins and disease protection.

Better post this before the computer here at the library terminates me.

Thanks again for giving me more things to think about. They will be fun to think about when I walk back home and start repotting all those little hellebores.

Best,
Elizabeth

Hi Elizabeth,
Very nice to hear from you. Sorry to disagree but I get far more fungal problems with red onions than white - especially neck rot and storage moulds. The usual hellebore received wisdom is that yellows are more susceptible than any other colour due to their odorus/cyclophyllus parentage. But I've not found it. I have virtually every colour and many species and still can see no pattern to fungal resistance either black spot, botrytis( the real pain!)or Cladosporium spp. This year due to the very mild wet winter followed by a sudden sub zero spell and then more humid clag I have fungal problems on every single colour. admittedly much easier to see on light colours but if you use a lens it's there on all, flowers, stems and leaves.
I have been to several shows and all the growers seem to be having the same problem. I do observe that lush growth resulting from forcing in polytunnels does seem to increase botrytis. Again not sure whether that is due to poorer resistance of plants or better conditions for fungi. I suspect like most things it's a combination of both.
The only plants that are doing well are the argutifolius and sternii - boots and tough spring to mind. Thibetanus is doing better than most (originates in a damp rain forest environment?)
Have you ever thought about the genetics of the different kinds of doubles i.e. true doubles where the petals/tepals double and nectaries are intact and the semi-double (most common form) where the nectaries have become petaloid. Must be different mechanisms?
I am carrying out some log term line "breeeding" experiments in this area but not mathematically. Can't see the point. What I am interested in is the mechanism and the interactions of genes - is there any epistasis?
Keep posting or contact me direct.
Mike

Hello all ,

Mike , always fine to meet a new responder .

The good news is that I could reach my Website as I could find a new provider .
With high speed , I've made some small changes and I added also a new pic of my white H x hybridus , the one that was blooming out of date ( July ) , but who is now making white petaloid nectaries . The plant must have 5 years .
It might be interesting to remember that this plant was highly stressed during the year . ( There are no doubles in my garden ! )

One of my observations with Rosa seedlings is that the first year they give mostly single flowers and from the second year on they double more . Also the number of petals depends on climatic conditions .

About the spots a lot of hypotheses can be made but is it possible to make crosses ( done by average gardeners ) to make one of these hypotheses harder ? Same question for other traits .
By browsing for "flower colo(u)r" , it was difficult to find good articles , mostly too simplified and wrong or too complicated . So I'm still doubting if the spots are of anthocyanin origin , yes everybody says so , but is this certain for hellebore ? or is it more complicated ?

To set up tests it is never bad to have some formation in mathematics , it helps a lot . Counting the results is not very difficult . The only problem I see is how to compare the results with those of other breeder .
If we could set up a dynamic data base , it must be possible to count the results of small tests together , I think .
If this exists , we may grab or use it .
If not we must think about two problems ; the user friendly layout solving the comparison problem ....and the script .

Johan

Here is a link that might be useful: fynwerk

Hi Johan,
looked at the link to fynwerk. Many of my singles show some degree of petaloid development of the nectaries without ever becomind a true semi-double(anemone). Some plants have single, semi double and double on the same plant. Usually the earlier flowers are more double and the later more single. I even have flowers of which half of one flower is double and the other semi-double! Have you looked at "hellebores" by Marlene Ahlburg. She seems to share some of my feelings about the non Mendelian complexity of hellebore genetics.
I have also line bred for 3 generations now a "true" double in which the tepals are doubled but the nectaries are intact. This produces a wonderful form as the "petals" are much flatter and less wispy than the usual nectary converted "petal" double forms.
Happy to share numbers but I still feel Mendelian maths won't provide the answers.
My hellebores are in flower from September to June in these mild global warming times.
Mike

Hi,

May be sticking my neck out, as I know nothing about Hellebore genetics, but found this thread of great interest. Basically, I have been a hobby grower and breeder of Hemerocallis (daylily) and am now getting interested in Saintpaulia (African Violets) but some of the facts from these come to mind as I read this thread.

First on doubling.

Hemerocallis is a monocot with 3 types of doubling.
A normal flower has 3 sepals, 3 petals, 2 whorls of 3 stamens each, and a tripartite ovary/style.

Type 1 double , the outer whorl of 3 stamens become petals
Type 2 double , Both whorls of stamens become petals
Type 3 double , either the petal whorl or the first whorl of stamens becomes converted to an complete apical bud, and this continues repeating giving a many petaled, generally sterile flower.

These are all genetically controlled by different genes, analogous to the ones described for doubling in arabidopsis (sorry, cant remember the reference right now). These appear to be controlled by supressor genes rather than active forward acting genes.

Spotting

In Saintpaulia there is a form called 'fantasy'. This is the appearance of random spots of one color on a field of another color. This is a dominate single gene, which apparently acts to randomly turn-on or turn-off one of the genes in the anthocyanine synthesis chain. What controls this gene is uncertain, but a single plant can vary somewhat from bloom to bloom, and vegetatively propagated plants from the same source can have any amount of activation from completely unspotted to completely the color of the spots.

Ignore this if it seems of no interest, just posted on a whim

George

Hello all ,

Thanks for the information .
George there is no problem to look how it works with other plants .
Myself , I started to learn about roses ( very difficult ) and hosta ( plastids ) and so on .
For the moment I go on with my tests as I can work with the F2 seedlings or second generation . They will flower this year and as I hope confirm my hypothesis , if not I'll tell everybody that the bees disturbed my statistics :-)

johan

Dear George:

Thanks for posting. All experiments are interesting and together we may make great contributions to plant genetics - or at least stretch our brains.

Dear Johan:

The old bees distorted my data ploy! Oh, no!