Rose Continuous Flowering Regulator

I have been emailed about the following Plant and Animal Genome XX Conference publication

The Continuous Flowering Regulator, RoKSN, Is Induced by Gibberellins to Modulate Flowering in Rose

Marie Randoux, Agrocampus Ouest, Institut de Recherche en Horticulture et Semence (IRHS); Julien Jeauffre, INRA, Institut de Recherche en Horticulture et Semence (IRHS); Tatiana Thouroude, INRA, Institut de Recherche en Horticulture et Semence (IRHS); Marjorie Juchaux, Université d’Angers, Institut de Recherche en Horticulture et Semence (IRHS); Jean-Michel Daviere, CNRS, IBMP; Amelia Gaston, INRA, UREF; Beatrice Denoyes, INRA, UREF; Hikaru Iwata, INRA, Institut de Recherche en Horticulture et Semence (IRHS); Laurence Hibrand Saint-Oyant, INRA, Institut de Recherche en Horticulture et Semence (IRHS); Fabrice Foucher, INRA, Institut de Recherche en Horticulture et Semence (IRHS)

I know the last two contributors, asked for communication and was allowed to post the link below to another publication about this subject:

Here is summary

Flowering is a key event in plant life, and is finely tuned by environmental and endogenous signals to adapt to different environments. In horticulture, continuous flowering (CF) is a popular trait introduced in a wide range of cultivated varieties. It played an essential role in the tremendous success of modern roses and woodland strawberries in gardens. CF genotypes flower during all favourable seasons, whereas once-flowering (OF) genotypes only flower in spring. Here we show that in rose and strawberry continuous flowering is controlled by orthologous genes of the TERMINAL FLOWER 1 (TFL1) family. In rose, six independent pairs of CF/OF mutants differ in the presence of a retrotransposon in the second intron of the TFL1 homologue. Because of an insertion of the retrotransposon, transcription of the gene is blocked in CF roses and the absence of the floral repressor provokes continuous blooming. In OF-climbing mutants, the retrotransposon has recombined to give an allele bearing only the long terminal repeat element, thus restoring a functional allele. In OF roses, seasonal regulation of the TFL1 homologue may explain the seasonal flowering, with low expression in spring to allow the first bloom. In woodland strawberry, Fragaria vesca, a 2-bp deletion in the coding region of the TFL1 homologue introduces a frame shift and is responsible for CF behaviour. A diversity analysis has revealed that this deletion is always associated with the CF phenotype. Our results demonstrate a new role of TFL1 in perennial plants in maintaining vegetative growth and modifying flowering seasonality.

and link

http://onlinelibrary.wiley.com/doi/10.1111/j.1365-313X.2011.04776.x/abstract

For those wanting more I am offered to ask for a PDF file. Let me know.

Hi Pierre!!

This is fantastic information and parallels and takes further that of Roberts et al. from 1999 and others. Roberts et al. suggested a Gibberellic acid gene with low expression in spring allowing for one time bloomers to bloom and then it gets ramped up to prevent rebloom and rebloomers have low expression at all times. It would be interesting if the authors of the recent paper addresses a possible connection with the gibberellic acid pathway. Roberts et al. looked at things from a perspective of what was different between a one time blooming rose and a repeat blooming sport of it and discovered some differences in GA. These other authors looked at the DNA sequence level. It would be great to make the link between the two with the biochemistry and pathways in between being better understood. With a strong understanding of these genes and sequence now, I bet it could be possible to develop repeat blooming roses of our favorite one time bloomers. With the major gene that governs repeat bloom here now better understood, one can use the sequence of the one time bloom allele and insert an antisense copy of it into the one time blooming rose. This is just the gene in reverse. The one time blooming rose allele and the inserted gene get expressed. The messenger RNA’s then are complementary to each other and bind. The one time bloom allele therefore cannot be efficiently transcribed. Theoretically we could get reblooming forms of our favorite species and one time blooming hybrids!!!

Oops I mean the one time bloom allele cannot be efficiently translated, not transcribed.

David, you may be right that it affects transcription or if it should happen to produce an RNAi effect. Very hard to predict exactly what will work or not, but things along those lines are certainly worth trying. The challenge will be to get the genes transferred into the relevant cells. You probably need to be culturing some rose parts and making somatic embryos. that’s a real challenge to do for any given CV.

David

Yes it is a new step discovering genetical modus operandi. Roberts et al. from 1999 and many others are among the numerous references of the copyrighted paper I saw.

And yes it allows considering future GM Continuous Flowering versions of actual Once Flowering vars or species. If it is all in a single gene…

Exploration of gene expression in different genome environments and/or other version of CF gene and/or looking for possible modifiers comes to mind.

It is plant architecture that one can consider manipulation.

I have been promised a here publishable reformated paper asap as well as a phone meeting with the team leader Fabrice Foucher about on going and future researches.

I got from Fabrice Foucher a long publishable PDF he reformated version of:

The TFL1 homologue KSN is a regulator of continuous flowering in rose and strawberry

Hikaru Iwata1,5*, Amèlia Gaston2*, Arnaud Remay3, Tatiana Thouroude3, Julien Jeauffre3, Koji Kawamura3,6, Laurence Hibrand-Saint Oyant3, Takashi Araki4, Béatrice Denoyes2 and Fabrice Foucher3,7

Apparently PDF file attachment is not possible here so I will send it by email to the asking ones.

Below are parts I selected:

SUMMARY

Flowering is a key event in plant life and is finely tuned by environmental and endogenous signals to adapt to different environments. In horticulture, continuous flowering (CF) is a popular trait introduced in a wide range of cultivated varieties. It played an essential role in the tremendous success of modern roses and woodland strawberries in gardens. CF genotypes flower during all favourable seasons, while once-flowering (OF) genotypes flower only in spring. Here we show that in rose and strawberry continuous flowering is controlled by orthologous genes of the TERMINAL FLOWER 1 (TFL1) family. In rose, six independent pairs of CF/OF mutants differ in the presence of a retrotransposon in the second intron of the TFL1 homologue. Due to insertion of the retrotransposon, transcription of the gene is blocked in CF roses and the absence of the floral repressor provokes continuous blooming. In OF- climbing mutants, the retrotransposon has recombined to give an allele bearing only the long terminal repeat element, thus restoring a functional allele. In OF roses, seasonal regulation of the TFL1 homologue may explain the seasonal flowering, with low expression in spring to allow the first bloom. In woodland strawberry, F. vesca, a 2 bp deletion in the coding region of the TFL1 homologue introduces a frame shift and is responsible for CF behaviour. A diversity analysis has revealed that this deletion is always associated with the CF phenotype. Our results demonstrate a new role of TFL1 in perennial plants in maintaining vegetative growth and modifying flowering seasonality.

A floral repressor regulates continuous flowering in rose and woodland strawberry

In rose, we demonstrated that the continuous flowering phenotype was explained by the insertion of a copia-like retrotransposon in the second intron of RoKSN, i.e. a TERMINAL FLOWER 1 homologue. Insertion of the retrotransposon modified RNA maturation. The second intron was not spliced out, and full length RoKSN RNA was not accumulated (Figure 4). In six independent vegetative mutants, climbing mutant roses differed from CF roses in the size of the insertion, probably due to recombination between LTR elements (Figure 3). After retrotransposon recombination, only the solo LTR element remained and the second intron was spliced out, thus restoring a functional RoKSN allele (Figure 4).

In rose, continuous flowerinf CF was thought to arise from a mutation in wild Chinese rose and then introduced in old Chinese cultivated roses (Hurst 1941, Ogisu 1996). We hypothesize that the mutation may be due to the insertion of a retrotransposon into the RoKSN locus in wild R. chinensis spontanea, as the retrotransposon was absent in the RoKSN locus in this wild species (Figure 3b), i.e. an once flowering OF rose that is supposed to be an ancestor of the original CF roses (Martyn 2005).

Pierre, thanks for sending this paper. It is very enlightening and should be on every rose breeder’s must-read list.

David, it looks to me like your conjectures over converting once bloomers into continuous bloomers will soon become reality now that there are specific dna targets and all you need to do, apparently, is disable their transcription. My guess is that they are involved in the ABA pathway rather than GA but time will tell. It wouldn’t surprise me if they already know which pathway is affected but are saving it for another paper.

From our perspective as breeders, knowing for sure that we are dealing with a gene that acts by knocking out transcription gives us confidence in assuming that intermediate levels of flowering are due to gene dosage effects. We can probably make reliable estimates about the zygosity for a lot of our breeders based on their flowering frequency and plan our crosses more wisely as a result.

One very important lesson that I derive from this paper is not to use climbing sports in a breeding program where remontancy is a goal.

Here is a report of what I understood and questions from a long phone exchange with Fabrice Foucher that quite friendly tried to explain things to the non trained in molecular genetics person I am.

First in simple words published findings are not revolutionary. Rather other (very important) steps that will allow further steps to better knowledge.

Not contrary to previous findings such as the florigene hypothesis.

Rose flowering is triggered by a promotor whose action is in our cultivated roses ancestors inhibited by a repressor. If both are expressed once flowering occur. If repressor is disabled then there is continuous flowering.

One inference is that all flowering plants may be supposed to have similar gene(s) triggering flower production.

Repressor disablement is known to originate from the chinese roses but variations were noticed by the publishing team (i.e. among climbing sports) and further work is planed to explore this at genus scale and probably more looking for alleles.

That there are variations among climbing sports repressor genes implies that the mutation is reversible and not so rare and eventually different ones occurred.

Thank you Pierre. That triggers the thought that perhaps the repressor can be like a dimmer switch, where instead of an absolute on or off situation, perhaps the weather allows it to respond by permitting flowering to continue as long as the appropriate conditions are present. This would be expressed when roses such as Banksiae continue flowering along the coast here for many months, sometimes up to eight or nine, before shutting off due to “hot” weather. In the same climate, O’Neal Blueberry continues flowering and producing berries. One year I know of, that continued nearly ten months when we had a very long, mild spring/summer. Nigel Hawthorne, which flowered just a few weeks in my mid desert garden, flowered four months in that coastal environment until someone bought it. I know there are many others which hit the threshold of what they require to flower, produce blooms for their appropriate period, then stop no matter what the weather does.

That species behave differently at some place out of their native habitat is not surprising. And banksiae are a good example.

Here french Riviera with a climate panel not unlike californian one the quite common banksiae flowering season is allways short and very early.Just like in all other places but some in California.

There is something there that longer inhibit banksiae flower repressor and this hints to a different one certainly worth looking at.

Years ago I reported about an anomaly: Chinas x banksiae F1 being all recurrent and never climbing just like further generations. An information I shared with Fabrice Foucher.

Pierre, here in the inland Southern California areas, which are more savannah-like, Banksiae flowers for maybe a few weeks. Closer to the coast, where the “hot” weather holds off for many more months, they flower repeatedly. Apparently, it’s the heat which triggers the flower repressor genes in them.

The farther inland you get from the ocean, the more extreme the temperature changes, both from night to day as well as month of the year. Along the coast, in many places, the most extreme the temperature differences between night and day is usually within twenty-five degrees. Often, it’s quite a bit less than that. Those more “spring-like” temperatures and light levels can continue from “winter” all the way through late August to September, when it begins to really heat up for a month or two.

To contrast, one mountain range inland, as well as higher up the mountains from the coastal areas, there can often be up to a fifty degree increase in temperature, with much less fog to soften the light intensities. Currently, the temperature difference between the coastal areas and where I am is between ten and fifteen degrees for day-time highs. By June, this can easily stretch to twenty and more daily. Nights are usually ten degrees warmer here during summer, and up to twenty degrees colder in winter.

Along the coastal areas, Banksiae can begin flowering in January and February and I’ve seen it continue until well in to July. Mine here in Encino, just over the top of the mountains from the areas where they can flower for six months and longer, began nearly two months ago and are still loaded with buds. The double yellow out front caps a large stand of Golden Bamboo, growing up through it and using it for support. This one receives sun all day long, with much reflected heat and light off the white styro foam roof. The bamboo causes it water stress as well as transpiring tremendous levels of moisture underneath it and through its mass. This one can spasm flowers just about any time of the year and it is nearly constantly mildewed.

Beneath and below the deck on the south-western corner of the house in the rear is a huge double white. It flowers later than the double yellow which is just across the deck, garage and apartment from it. This one has no other plants close enough to it to produce the humidity to induce the mildew, and it is often as dry as the one in the bamboo as the only irrigation is rainfall and when I happen to feel I want to hand water it with the hose. This dry corner is significantly hotter than where the double yellow is, but receives only two-thirds the direct sun the double yellow receives. Both grow rampantly, but the white begins flowering weeks later, having been in bloom now for about a month. The lutescens is still potted in a twenty gallon squat and hasn’t begun setting buds yet. It also receives filtered sun through a California Black Walnut until mid afternoon, when it receives full, direct sun from the due west exposure.

Vina Banks is planted out to the west of the double white on the hill and has popped flowers much of the winter and still has some today. There is a Purezza in the front walled garden I’m hoping will spill over the front wall. It doesn’t have as much reflected heat as there is climbing fig on the wall which softens the reflected heat. This one flowers mainly when our temperatures are well into the eighties F for extended periods. The canned Purezza (still waiting to find a home in back), is shaded by the Walnut and has thrown a few flowers but nothing like what I expect when it gets hot. Right now, we’re in the mid seventies, but have had several spikes to the mid eighties in the past three weeks.

By September, there can easily be up to a fifty degree temperature difference between the coast and right here, with several degrees even hotter as you travel further inland than I am. I’ve left work right on the ocean on Labor Day (early September) where it was very foggy and sixty degrees F. Within five miles, it was already in the mid nineties F. By the time I had driven to the center of the valley I live in now, it was 115 F. Fifteen miles further, at home, it was 119 F. Even just five miles up the moutain in The Highlands, in the same postal zip code, it can be fifty degrees hotter with no fog, while the bottom of the mountain is cold, damp and extremely foggy. This is considered “normal” for the area.

In that fog belt is where the Banksiaes continue flowering well into the year, unless the weather turns hotter, earlier, when they cease flowering. Mine begin weeks later and are usually finished when the heat begins, months earlier than along the coast. In the Santa Clarita Valley, the next valley inland from here, they are finished months earlier than even here, because the heat has usually already started intensifying and remaining hot.

I’ve seen the same performance from the hybrid Hulthemias, Nigel Hawthorne and Euphrates; R. Arkansana “Peppermint” and O’Neal Blueberry to remember a few. On the edge of the fog belt, I had a client who planted many O’Neal Blueberry because her four year old daughter loved them. Those bushes began flowering in January (as mine are here, now) and continued to flower and fruit until mid September, when her garden got “hot”. The roses came from Ashdown and were budded on R. multiflora, canned and sold where I worked. The same varieties from the same source, performed here as expected, as does the O’Neal Blueberry. Though, one block from the ocean where it was colder, extremely damp and heavily foggy much of the day and where the temperatures remained in the low to mid sixty degree range, the three roses mentioned above and the Banksiaes, began flowering earlier and continued until at least late August when it heated up that year. Nigel Hawthorne continued one year until mid September, as did the Banksiaes, as it got “hot” there later.

That all leads me to feel it is temperature induced, whether it is just the heat level, or the extremes between high and low temperatures, I don’t know, but “hot” seems to turn off the flowering in the traditional Banksiae, the roses mentioned earlier and the blueberry. Eucalyptus begins flowering in the fog zone weeks earlier than here and continues a good month later, also depending upon how hot it gets, how early it happens. Vina Banks and Purezza require the heat to begin and continue flowering. In the chillier zones, they can begin flowering once there is sufficient heat, but stop until the much hotter weather begins. Once, I’ve been told the Purezza out grew that performance, beginning to flower when expected and continuing to repeat, even after traditional Banksiae have stopped. Inland, where it’s hotter, Purezza begins usually when the traditional Banksiaes stop and continue flowering heavily as long as the weather remains at least in the high eighties F. Kim

At these temperatures the dominant mechanism preventing flowering - and plant growth - is photorespiration.

Photorespiration is the opposite of photosynthesis whereby carbon is lost to the atmosphere. At nominal temperatures photorespiration rates are much less than photosynthesis so a plant can accumuilate carbon and make starches with it. However, the rate of photorespiration increases with temperature such that by mid-90°s F more carbon is lost than is accumulated and growth stops.

This has other implications for breeders as well. Programs such as Earth Kind that select for plants that can tolerate high temperature conditions should consider that photorespiration presents a fundamental biochemical limit to the ability of a plant to grow.

Thanks Kim

Difference is that here the sea is hotter and coastal fogs if not unknown are rare, short lasting and quite localized so that they are not effective at lowering temps.

In Europe actually we are undergoing a cold spell. Out of sun ice does not melt over day. These nights in the twenties are excellent for future germinations.

Rose optimum temperature is not high. Once I was in Rio de Janeiro and its nice Botanical Garden. There is a large area devoted to a rose collection that was quite pathetic. Just surviving small baren sticks. Only Flower Carpet had more leaves hanging on but so desease ridden it was very difficult to recognise.

Roses adapted to hotter parts such as hulthemia or Hesperhodos have small leaves and habilities to survive not grow in heat and drough.

These findings about roses all having genes for recurrence that are expressed when the repressor are disabled explains how some chinese roses were recurrent.

And here we could better understand leaning on ancient chinese roses history. A very ancient history that began collecting wild roses for centuries and over a long long time lapse (tens of centuries) selecting among seedlings the more garden worthy ones that were found to be double flowered, recurrent or dwarfer. Culminating in Parks Yellow all modern HTs are so heavily built on.

The chinese roses are a very mixed bag. Some are close to species: i.e. banksiae are true breeding with nothing in beetween the single and quite double yellow and white. Never creamier or semi double.

There are also many vars with the gigantea-chinensis look.

But to name but two Slaters or Parsons have a partly obscure ancestry.

To be continued…

Did a quick online search on the effects of temperature and gibberilins, and found this article on dwarf apples of potential interest:

http://onlinelibrary.wiley.com/doi/10.1111/j.1399-3054.1992.tb02167.x/abstract

It didn’t say anything about flowering or fruit-set with regards to such, but the presence of GA’s and elongation of growth would seem to validate Kim’s observations.

And Don, while your comment on breeding with climbing sports makes sense on the face of it, I would be interested in knowing if that argument is born out in other folks’ practical experiences. I’m not sure how different the genome is in a sport exhibiting a differing phenome such as climbing and the consequential reduction in flowering. The “on-off” switch might be more nuanced than that.

It is interesting to note the sales claims of some roses’ climbing sports which purportedly do not exhibit a reduction of remontancy. I wonder what the implications are for using these roses in breeding. Why might one switch occur and not the other, if, as presumed, GA levels change in these sports?

Philip, those not exhibiting reductions in remontancy are historically recorded. We not only have the Cecile Brunner/Spray Cecile/Cl. Cecile Brunner with varying plant sizes and remontancy levels, but also the full-on climber which eats barns and some rebloom while others don’t. New Dawn has exhibited the same variability, not only obviously climate manipulated but from plant to plant in the same climate as if it only partially mutated to repeat flowering, which is the case, I believe, with Cecile Brunner and her variants.

Old ARS annuals reported various mutations of the same HTs to climbing forms, also with varying levels of flowering. Suggestions were frequently made to make sure the stock obtained came from the particular sport from particular nurseries. I’ve read reports from various nurserymen through the old annuals where they stated they’d obtained “better stock” of climbing sports, from the older HTs to Iceberg, to insure what they offered had the highest level of rebloom. There have been numerous commments about Blaze Improved throughout the decades, also, and its ability to perform as original, once-flowering version to more fully repeating type. Of course, some of those could also be climate induced.

It’s been interesting following breeding from the various climbing mutations. Some breed climbers, while many others don’t. Of course hybrids of New Dawn should be climbers, as it mutated from a climbing seedling, so climbing tendancy exists at the sexual level. Encountering climbing seedlings resulting from bush types crossed with climbing mutations of bush types seem to demonstrate the depth of the mutation. I remember correspondence with Dr. Michael Dykstra way back when he was breeding with striped sports of HTs in hopes of creating new striped HTs. I suggested to him he should use Ralph’s striped minis as they demonstrated the abiltiy to generate striped seedlings, while non of the mutated stripes I’d ever raised seedlings of ever demonstrated any variegation in petal colors. Ralph stated he’d selected Ferdinand Pichard to use because it was the only striped rose he had access to which had no stated parentage (read not stated to be a mutation), hence he felt it possible it was seedling raised and possessed the ability to pass on stripes.

A further facet of the Banksiae thread…my across the street neighbor has a Banksiae lutea next to his house, receiving southern exposure, reflected/radiated heat from his house wall, fronted by lawn. His plant is large, also, and hasn’t begun flowering yet, while mine is loaded with bloom. They receive the same direction and intensity of direct sun, for nearly the same period, while mine probably gets more light, and perhaps a little more heat, with more humidity than his due to the bamboo and ultra white, quartz surfaced, styrofoam roof. I know the light level is tremendously greater off the roof, while the actual heat may not be as great as that reflected and radiated from the earth tone cement wall of his house. They are at the same elevation. My plant’s trunks and roots are continuously cool, growing under the bamboo and through its mass into the sun. His heats up from the base up. I’m not entirely sure how to interpret the data, but it is interesting. I believe his receives nitrogen feeding from what’s put on the lawn. Mine only receives “organic” fertilizing from the dense bamboo litter falling into the clump from above. His definitely receives much more water from lawn irrigation than mine does.

The “on-off” switch might be more nuanced than that.

Hi Philip,

At first I thought that Foucher’s data says otherwise. They examined seven pairs of roses and their climbing sports and in every case the mechanism was a single switch event: “In OF climbing mutants, the retrotransposon has recombined to give an allele bearing only the long terminal repeat element, thus restoring a functional allele.”

Moreover, the effect of a single transposon held across ploidy levels for the three pairs they tested for which we have ploidy data, Old Blush (diploid), Iceberg (triploid) and Peace (tetraploid).

Still, as Kim points out, we have a pretty reliable catalog of mutant pairs that don’t fit this profile. It makes you wonder whether they accidentally cherry-picked cultivars all having a common ancestor bearing the original transposon and missed others that had a different mutation. Maybe they should treat Kim’s list as a menu for further investigation.

Can folks here explain how true climber roses (ie. not sported climbers) tend to behave in breeding.

In particular I am interested to know with what frequency that “sexually acquired” climbing habit is inherited in rose breeding.

George, you have asked part of my question which I have been thinking for a couple of days. Can I go one thing/step further. I am wondering if roses came as climbers or sported from bushes. I know, go and read from somewhere the history of roses, it will tell me. But in short did someone “find” climbing roses.

This is, as they say, a very thorny subject. Some species of rose like gigantea look like what we’d all agree is a climber. But what of multiflora? Just a big bush ? even if it is 15 ft up in a tree like some in PA that I know of. On the other hand High Noon when I saw it in the Pacific Northwest was unquestionably climbing massively and blooming like crazy. It was made that way, not as a sport of anything else. I grew it in a 5 gal bucket for years and got regular blooming on an excessively vigorous plant a lot like Captain Thomas which is in its pedigree.

My climbing sport of Crimson Glory gave (self seedlings) only climbing types that bloomed basically once a year, except occasionally in fall after severe summer stress and fall rains. But I know that in Philly, there was a climbing Peace back in the 60s that bloomed all the time.

New Dawn is a hard example because it’s triploid. Van Fleet (parent) blooms only once a year. But we don’t yet know how many copies of what genes it carries in the flowering locus series. Parentage is not real clear. Self seedlings of N.D. generally don’t bloom first year so get discarded ( I never got 1 in 50). But lots of crosses with mini pollen have given early, repeat blooming. Number of leaves before a bloom is a variable with basal breaks on those. One I’m focused on right now fro breeding makes tall, late flowering basal breaks with a single flower at the end, like N.D. IN spring the laterals bloom as expected. N.D. itself sends canes with a terminal flower and not all that many leaves, maybe 10-15, just very long internodes. Then in the same year, if the season lasts long enough, secondaries bloom on shorter stalks. This is true in my climate, maybe not elsewhere. My N.D. was propagated from flowering laterals at the approach of winter so was selected for good rebloom.

Early in their breeding the Brownells had a number of what they called pillar roses. The distinguishing characteristic is to make a long cane with a terminal flower. Then branching laterals. Rhode Island Red or Golden Arctic are examples I’ve grown a lot. Some famous parents in this line were clearly what almost anyone would call a climber, or rambler. Obviously there is a blending of behaviors here. I think we need to count the leaves per cane, not just length. Last summer my peachy creeper had 42 leaves below the flower on a 2 ft or so cane while N.D. had a considerably longer cane with just 14 leaves.

The HP/HT Frau Karl Druschki usually shoots out some very long indeterminate canes here toward midsummer… Most of the spring bloom comes as laterals from these. But if the long canes are pegged down they may break forth flowering laterals later in the fall. And some years, like last summer, it was scarcely out of bloom all season, with one or two at a time on canes of all lengths.

The authors did hedge their bets a little by stating there may be secondary effects. Yes, I think they only looked at a few HT/Fl shrub/climber pairs. But we do know that for a tetraploid HT to turn climber it takes only one mutation. For instance Crimson Glory, Peace etc. This says the gene is dominant. But I’d expect with two copies the dominance would be greater if quantitative genetics apply. So how easily the climbing type can rebloom will depend on modifiers like climate or the levels of expression of various hormone genes. That’s why I say we need to count the leaves below the bloom.

Thanks, Larry! That was interesting! You’re saying, then, it shouldn’t be a “climber” if there are higher numbers of leaves below the flower? I’m trying to understand and interpret what I should look for and what it should mean. Thanks!