Inheritance of Winter Dormancy

Many Teas and Chinas will grow whenever the ambient temperature is warm enough. Rosa multiflora, on the other hand, goes dormant in the fall, and stays “asleep” for some time. I would expect the progeny of a cross between Multiflora and a China or Tea to segregate in the F2 for winter dormancy, but I have found no published study.

Has anyone observed this sort of segregation in other crosses of winter-dormant and non-winter-dormant varieties? Are there any published reports I’ve missed?

I have searched for information on the inheritance of winter dormancy, but most of the reports I’ve found have dealt with efforts to reduce the chilling requirements of fruit trees (apples, peaches, apricots, etc.). In breeding for hardiness in roses, it would be useful to increase the chilling requirement to delay sprouting in the Spring, as well as to select for earlier dormancy so the plants will be prepared for early frosts in the Fall.

Some roses are surprisingly hardy. ‘Gloire de Dijon’ tolerates winter better than other Teas because it has a proper dormancy, presumably with a chilling requirement that prevents it from “waking up” during warm spells during the winter. ‘Marechal Niel’ reportedly survived a winter in Oklahoma that killed various other roses that had been bred for the far north by Brownell and Horvath. I doubt that ‘Marechal Niel’ has a proper dormancy, but it does love heat – a fact that suggests it would be unwilling to grow at low temperatures.

Some roses of the far north apparently lack dormancy. They don’t need it because the continuous cold prevents them from growing. The octoploid R. acicularis suffered winter-kill at Ann Arbor, MI. And seeds of R. blanda collected in northern Michigan segregated for tenderness and hardiness when raised at Ann Arbor, 300 miles south of their original home. No doubt these “tender” seedlings would have done well back home where the continuous cold restrained their growth.

It would be interesting to cross Marechal Niel with Gloire de Dijon in order to combine the heat-loving property of the former with the dormancy of the latter. Such a cross would be more likely to give hardy offspring than any cross involving the octoploid R. acicularis or one of the R. blanda seedlings that was tender at Ann Arbor.

One more thing: I have read that the descendants of ‘Rival de Pestum’ are hardier than most other Chinas and Teas. Does anyone have experience with any of these (Comtesse du Cayla, Mme. Laurette Messimy, Madame Eugène Résal)? Does ‘Rival de Pestum’ or its offspring go dormant in Autumn?

And some notes on this subject:

The Gardener’s Monthly 2(3): 69 (Mar. 1860)
HARDINESS OF CHINA ROSES
CHARLES G. PAGE, WASHINGTON, D. C.
There is probably no variety of rose that will endure a temperature of zero Fahr. upon unripened branches, and expanded leaf buds; and there are probably very few roses that will not endure this temperature provided the wood has been fully ripened and the buds are all dormant and the sap quiet. The Gloire de Dijon is an excellent illustration. It belongs to a tender family, but is perfectly hardy here. Its hardiness is not, however, entirely intrinsic, but depends upon its habit of growth. Unlike Teas and Noisettes generally, it stops growing in the fall, and is not apt to be quickened again till the spring. It prepares for winter like a Remontant, and has proved itself here more hardy than the majority of Remontants. In that rigorous winter of 1855-6, it stood better than La Reine, Madame Laffay, Wm. Griffith, and others. This winter has been thus far very destructive to Teas and Noisettes, but the Dijon is unharmed.
http://bulbnrose.x10.mx/Roses/breeding/PageHardyBengals1860.html

The Rose Garden – Division 2, Page 124 (1848)
William Paul
The Gloire de Rosomene suffers from severe frost: the progeny is hardy.

Bot. Gazette 96(2): 197-251 (1934)
EXPERIMENTAL DATA FOR A REVISION OF THE NORTH AMERICAN WILD ROSES
BY EILENE WHITEHEAD ERLANSON
p. 207
R. acicularis from Alaska (2n = 56) responds so rapidly to a rise of temperature in spring that the flower buds are almost always completely frost-killed in April in southern Michigan. The hexaploid requires more or prolonged warmth and always flowers profusely.

Seedlings of R. blanda from northern Michigan segregated into tender and hardy individuals at Ann Arbor, 300 miles south of their natural habitat.
http://bulbnrose.x10.mx/Roses/breeding/Erlanson/ErlansonRevision1934.pdf

New Phytol. 37:72-81. (1938)
PHYLOGENY AND POLYPLOIDY IN ROSA
By EILEEN W. ERLANSON, D.Sc.
pp. 77-78
A plant of octoploid R. acicularis from Alaska was transferred to Ann Arbor, Michigan, where it leafed out soon after the first April thaw and was frequently badly damaged by frost in May. In some seasons all the floral primordial tissue was destroyed and no flowers were produced. The alternating thaws and frosts which are a feature of continental climates between 40 and 50° of latitude North, may militate against the spread southwards of octoploid races in Rosa. The Alaskan octoploid thrived at Pasadena (Erlanson, 1934) in southern California, and came into flower early in February, 3 1/2 weeks before the hexaploid type.
http://bulbnrose.x10.mx/Roses/breeding/Erlanson/ErlansonPhylogeny1938.html

American Rose Annual p. 103 (1943)
Maurice H. Merrill
Normal, OK
In my garden the Brownell and the Horvath productions, bred for resistance to winter cold in northern latitudes, so far, with the exception of Mabell Stearns, have displayed a susceptibility to severe, and often fatal, winter injury. In contrast, the hardiest, least winter-harmed bushes I have today are Old Blush, a China which is close to a Tea, and a nameless waif I acquired on our farm, where it had been brought by the tenant’s wife who found it at a roadside filling station. Federation shows much more damage from this last winter than do three young Marechal Niel plants, not yet fully established. All this leads me to the none-too-profound suggestion that the qualities which make for hardiness in the long, severe northern winters, in which a rosebush can hibernate like a bear, may not facilitate survival in the open winters of the Upper South, particularly our western portion, punctuated with occasional periods of severe weather.
http://bulbnrose.x10.mx/Roses/Rose_Pictures/M/marechalniel.html

American Rose Annual 41: 118-122 (1956)
Difference In Resistance To Spring Freezes In Rose Varieties
Dr. H. R. Rosen
In general those varieties which have an inherent tendency to break their dormancy ahead of other varieties are apt to suffer more from late spring frosts and freezes. For example, Climbing Mme. Edouard Herriot, one of the most beautiful and hardier sports of a Hybrid Tea when it is dormant, having withstood temperatures below zero while unprotected in my own garden, has the habit of breaking its dormancy considerably ahead of such varieties as Paul’s Scarlet Climber, New Dawn, Bonfire, and Chevy Chase. It had much new growth and numerous blossom buds when the March freezes struck and although about twenty years old, it was completely killed. On the other hand, while the other varieties just mentioned suffered a loss of 50 to 90 per cent of their wood, all of them survived.

Another factor which tended to obscure the relationship of winter hardiness to resistance to spring freezes is rapidity of growth. Contrary to what might be expected, those varieties which made the most new growth prior to the freezes were the ones which were hurt the worst. Thus in the fungicidal test plots where two old standard varieties have been used for many years, Etoile de Hollande and Edith Nellie Perkins, the former suffered far worse than the latter although both broke their dormancy about the same time, as they usually do. Etoile de Hollande usually makes about twice the growth in a given length of time, as it did prior to the freezes. Out of 88 plants of this variety, 49 were completely killed and the remainder were set back to such an extent that they probably fell far below average in seasonal amount of growth and of blossom production, while of 96 plants of Edith Nellie Perkins, only 34 were lost and the growth and seasonal blossom production of the remainder appeared to have suffered very little. However, as no blossom counts were made this year because of loss and injury, the data are incomplete.
http://bulbnrose.x10.mx/Roses/breeding/Rosen_Spring.html

American Rose Annual 26: 111-115. (1941)
The Search for Total Hardiness
Georges Bugnet
With a much longer experience, on a larger scale, in stone-fruits breeding, I am led to believe that a plant, in order to withstand our climate, needs a very early ripening of its tissues. Winter-killing, apparently, is not caused by extreme cold but rather by a too early cold snap catching immature wood, like the 30° below we had in the first part of November last. Once, at dawn, on October 12, 1930, we had 16° below zero. The next day was rather warm. None of my hardy hybrids and no native tree or shrub suffered. I have often noticed that half-hardy plum or apple trees here, unhurt by December 1, passed unharmed through the rest of the winter no matter how intense the cold.

Native trees here, as a rule, put out their leaves around the middle of May and drop them near the end of September. Roses have also to be brought to an early ripening of their tissues.
http://bulbnrose.x10.mx/Roses/breeding/Bugnet.html

Karl,

Great posting. There’s a lot to read and absorb but my initial thought was would it make any sense to combine ‘Gloire de Dijon’ with the two hardy species you mentioned (R. acicularis and R. blanda) to add dormancy genes?

Rob,
I wouldn’t think so. I suspect that those two species are cold tolerant — they like growing in cooler weather (what choice do they have?). This is a problem with derivatives of R. wichuraiana and R. foetida, as well. If the crosses were grown near Ann Arbor or further south, they would likely start growing too soon in the Spring, even with dormancy. Jerabek (1975) wrote of yellow HTs, “As a class, winter hardiness is poor and I have discarded many varieties for this reason.” I think this is due to their habit of sprouting too early. I don’t know how heat-tolerant ‘Gloire de Dijon’ is, which would be an important consideration.
http://bulbnrose.x10.mx/Roses/breeding/JerabekYellows1975.html

I played with the idea of crossing ‘Golden Showers’, which is just about worthless in the Great Plains region because it stops growing in hot weather, with ‘Marechal Niel’, which lacks a proper dormancy but LOVES heat. A derivative of such a cross might be as hardy as GS, but as happy to grow and bloom in the summer as MN.

An approximation of this is ‘Diamond Jubilee’ [Marechal Niel x Feu Pernet-Ducher]. It blooms well in heat, as I learned in Lexington, KY during the hottest part of summer. However, I don’t know how far north it will grow successfully.
http://bulbnrose.x10.mx/Roses/Rose_Pictures/D/diamondjubilee.html

I forgot to mention that the ‘Orleans Rose’ group of Polyanthas seem to have a longer winter dormancy than the sport-offspring of ‘Tausendschon’ (all the Koster clan). I found this annoying when I lived in California, but now I’m thinking happier thoughts about ‘Orleans Rose’ and its kin.

I think this is related to the question of whether flowering is day-length dependent in repeat-flowering roses. I think that it is, for some CVs. For instance, Winter Sunset, a Griffith Buck rose has its best blooms in late fall, when treated exactly like other Buck roses that have already shut down. So in my interpretation, WS has less daylength dependence of flowering than the others do. It could also be total amount of sun affecting photosynthate production. A bit hard to sort out a true daylength signal from overall growth response to cooler shorter days. But it is still true that WS blooms better in those cooler shorter days than some other CVs.

Hip load makes a big difference too. My original Carefree Beauty (grafted) set hips and continued reblooming. It was in full sun. Now I have a mature own-root plant which tends to shut off flowering if all the flowers of the first crop are successfully pollinated. But it is in significant shade. With heavy feeding a smaller potted plan will produce some later blooms in the presence of hips, but I know it would bloom more abundantly if dead-headed.

Diverting sugars into osmolytes as a response to shortening days is key to tolerating cold by allowing supercooling. It is hard to maintain both full flowering and cold-resistance at the same time. They are responses to opposing signals. It is very challenging to design an experiment to sort out the daylength (phytochrome) vs energy production factors.

Thanks Karl for this article.
Like Rob Byrnes says this is a very difficult to digest at once.

Many of the Canadian Explorer Roses have been hybridized with Hybrid Teas and floribunda in their back ground.
The question I have for members in the central and south USA. How do the Explorer Roses perform in these Warmer climates?
These roses were bred for very cold areas, do they shut down in the heat of midsummer.
Chuckp

I would be happy to see more research into the effect of photoperiod on Rosa species and cultivars. There was a species (I forget which one) at the Heritage Rose Garden, San Jose, that dropped its leaves well in advance of the thousands of other specimens. Photoperiod response would be a plausible explanation for this, because I doubt that even the night-time temperatures were low enough to provoke dormancy.

As for ‘Winter Sunset’, you might also consider what sort of fatty acids that variety produces, since this can influence when a plant grows most happily (saturated FAs favor growth in heat, unsaturated are better for cooler temps.)
http://bulbnrose.x10.mx/Heredity/King/FattyAcidBiblio.html

This temperature preference is also expressed in seedlings. Risley (1958) studied the influence of various pollen parents on seedlings raised from ‘Skinner’s Rambler’. Unfortunately, he confused the matter by assuming that only seed dormancy was involved. It struck me as vary odd that a hardy pollen parent like ‘Persian Yellow’ would father offspring with a shorter dormancy, while ‘Diamond Jubilee’, a seedling of ‘Marechal Niel’, would contribute to a longer dormancy in its progeny. These results seemed backwards. But if seed-dormancy was not the main issue, the slower growth of DJ offspring struggling to emerge while still in cold storage would make sense.
http://bulbnrose.x10.mx/Roses/breeding/Risley.html

It was this realization that led me to search for more info on temperature preferences. Here’s some more info:
http://bulbnrose.x10.mx/Roses/breeding/Greeley1919/Greeley1919.html

How does ‘Winter Sunset’ compare with other Buck roses (and other hardy rose in general) with its sprouting in the Spring?

And while I’m at it, I should mention that soil can influence hardiness. Clay soil was traditionally regarded as “late” because plants grown in clay are slow to emerge because plants aren’t going to do much while their roots are trapped in cold, clammy clay. It is apparently for this reason that Rosa arvensis was found to be rare in the North of the UK, but where found it was in clay.
http://bulbnrose.x10.mx/Roses/breeding/Allen_R_Arvensis.html

I just found a couple of papers on the inheritance of chilling requirement in apricots.

ISHS Acta Horticulturae 622: XXVI International Horticultural Congress: Genetics and Breeding of Tree Fruits and Nuts
INHERITANCE OF CHILLING REQUIREMENT FOR DORMANCY COMPLETION IN APRICOT VEGETATIVE BUDS
R. Tzonev, A. Erez
Abstract: The pattern of vegetative bud dormancy was studied in apricot hybrid populations and the inheritance of chilling requirement evaluated. The study was carried out in Israel in a region with mild climate under natural conditions after two consecutive unusually warm winters. Vegetative bud breaking (BB) was estimated by evaluating 3 parameters: the level, the pattern, and the timing of BB on each of 1529 hybrids obtained from 44 crosses or open pollination. A most striking result was the high level of vegetative bud break registered especially during the first year of our two-year experiment, in progenies of all cultivars and selections including ones with high chilling requirements. On the other hand, a non-uniform pattern of vegetative development, reduced growth rate and a late bud breaking were characteristics found in both years in crosses with high chill parents. From the data obtained we concluded that the chilling requirement in apricot vegetative buds represents two distinct phenomena. The first is a genetically controlled switch of bud break, and the other phenomenon is the genetically control of vigor of the ensuing bud growth. The inheritance pattern of the first trait seems to be low chilling dominant while the second exhibits a non-dominant intermediate response between the parents. Both genes require exposure to chilling prior to activation but the first one responds to a lower level of chilling.

It is useful to note that bud growth and the ensuing bud growth seem to be inherited independently. Rosen (1956) reported that the roses he studied varied in the growth rate of new shoots in the Spring, as well as in the timing of bud break.
http://bulbnrose.x10.mx/Roses/breeding/Rosen_Spring.html

According to the other report, by Viti, et al., “Analysing the flower bud dormancy evolution of genotypes of the two progenies, a different distribution of character CR in relation to the cross combination was observed. From deep to the end of dormancy, the glutathione oxidised form GSSG was depleted and the active reduced form (GSH) increased, altering the GSH/GSSG ratio. Although, this behaviour was different in relation to the genotypes, glutathione is one of the putative key compounds inducing the resumption of flower bud growth.”

For those of us who don’t have the facilities for measuring GSH/GSSG ratio, it is helpful to know that a specific substance is involved (assuming this also applies roses). Thus, it should be possible to select directly for longer dormancy (and indirectly for increased GSSG).

Willows are far removed from roses, of course, but this report suggests places to look for differences in resistance to “winter kill” due to early frosts and freezes.

Lennartsson, M. 2003. Cold hardening and dehardening in Salix. Doctoral thesis. Silvestria 279.

“The variation in cold hardiness in Salix in the autumn was investigated using clones of different geographic origins. In late growing season, the variation was small and inversely related to a phenotypic variation in potential growth rate. When growth had stopped in response to the reduction in daylength, however, large differences in cold hardiness developed. Northern/continental clones started cold hardening up to two months earlier and showed up to three times higher inherent rates of cold hardening than the southern/maritime ones. The two components of cold hardening, the timing of onset and the inherent rate, seemed to be separately inherited traits, as judged from analyses of the prodigy [sic] of a crossing between an early-and-rapidly hardening clone and a late-and-slowly hardening one. This suggests that cold hardiness can be improved without adversely affecting growth by selecting for a late onset of cold hardening combined with a rapid rate. Also, in the early stages, cold hardening was more sensitive to low, non-freezing temperatures in the southern/maritime clones than in the northern/continental ones.”

This item fits with what Lennartsson (previous note) observed. However, it adds that what happens in Spring (late emergence, fast growth) mirrors what happens in Fall (early and rapid hardening).

Some Problems of Method p. 44-51 (1934) 1952
Means of Shortening the Vegetative Period of New Plant Varieties
Ivan Michurin

The ten new hybrid varieties of hardy grape—which stand the winter without any artificial protection—that I have produced in late years made it possible to extend the zone of cultivation of the grape another five hundred kilometres northward.

Besides winter-hardiness, what is needed of the grape for the advancement of its cultivation northward is later flowering, because of the spring morning frosts, and earlier ripening of the fruit, because of the early autumn frosts. All this together presents a difficult problem, namely, how to shorten the vegetative period of the grape.

Already in the 1900’s, while working on hybrid varieties of yellow cigarette tobacco, the Kommunarka early-ripening melon and hardy grape seedlings—the first to be produced in those days—I was agreeably surprised, when selecting seedlings that completed their vegetative development earlier than others, to find that some of the seedlings that had germinated from seed later than others, namely, at about the beginning of July, managed to complete their growth and mature even earlier than those that had germinated in the middle or beginning of May.

I made a note of this marked, and at the same time rather paradoxical, phenomenon, and in subsequent years I never failed to keep watch for similar manifestations in interspecific hybrids of other plants. It turned out that this phenomen is in most cases to be met with in hybrids from parents whose habitats were very far apart, and that, on the contrary, it is practically never encountered in simple seedlings or in hybrids from varieties of one and the same species coming from mutually close places of origin.
http://bulbnrose.x10.mx/Heredity/Michurin/MichurinGrapes1934.html

In addition to dormancy and preferred temperature for growth, hardiness (or resistance to “winterkill” involves a plant’s ability to protect itself from chilling injury by producing antioxidants such as ascorbate, glutathione and carotenoids in chloroplasts. Sugars have been thought to act as “antifreeze” in the cells, and this may be true. But glucose is also necessary to synthesize ascorbic acid (vitamin C) and ascorbates.

Chlorophyll fluorescence gives a measure of chilling injury, allowing breeders to assess resistance to chilling injury under controlled conditions.

Report of the Tomato Genetics Cooperative,
Volume 41 (July, 1991)
Physiology and genetics of chilling tolerance in tomato
Walker, M.A. and Smith, D.M.
http://tgc.ifas.ufl.edu/vol41/v41p62a.html

HortScience 35:184-186. (April 2000)
Assessing Chilling Tolerance in Roses Using Chlorophyll Fluorescence
by Nadia Hakam, Shahrokh Khanizadeh, Jennifer R. DeEll and Claude Richer .
http://www.google.com/url?q=http://www.researchgate.net/profile/Shahrokh_Khanizadeh/publication/52010518_Assessing_chilling_tolerance_in_roses_using_chlorophyll_fluorescence/links/0912f50eacc8759a46000000.pdf&sa=U&ei=GQ01VZXfKcecNqCHgcAB&ved=0CBQQFjAA&usg=AFQjCNEY28DXT_xE6aoeh6QMYEBgoBt_qQ

I think Karl is quoting a very interesting point in the penultimate post above, namely the early observation of Michurin about late-popping seeds giving short season plants. Note that this was emphasized as a phenomenon in wide crosses. So it suggests to me that my patience may be rewarded. Keeping the stratifying seeds for a year is a real pain but may be worth it. I will try to pay more attention to this as I go along. Last year I kept on planting out sprouts right through the fall, maintaining them indoors over the winter. I did get several interesting ones, now i need to see if they are better adapted than would be expected from their parental pedigrees. Some of the late ones were about the only germinations from particular crosses so it’s hard to know if this is a useful trait or a sign of a screwed up genetic combination.

Another related observation- winter survival of seeds. I found it rather hard to believe but for some CVs, hips that were exposed to the winter’s cold outside can give very high germination. That tells me that at least the embryos are able to get themselves into a state of cold tolerance such that they can survive down to 5 F or so. I think this might be a good head start of real cold hardiness. White out is the CV that comes to mind at the moment. A rugosa growing in Lincoln NB, (where it got considerably colder than 5 F) is another. Really not unreasonable in the real world but not what most breeders have selected for over the past century.

Larry,
The fact that Michurin observed this phenomenon in the first generation suggests that it may be a case of what I call “elective expression”, which seems to be more common “wide hybrids”: interspecific, intergeneric or between geographically isolated races.
http://bulbnrose.x10.mx/Heredity/King/Election-Variegation.html

For example, Karpechenko (1924) pollinated a Radish by various types of Cabbage, Kale and Brussel Sprouts. Some of the F1 hybrids had rough leaves similar to those of the radish. Some had smooth, waxy leaves like the cabbage parent. The remainder, the giant plants, mingled the characters of both parents.
http://bulbnrose.x10.mx/Heredity/Karpechenko/Karpechenko.html

Hurst (1900) reported similarly for Paphiopedilum hybrids.
http://bulbnrose.x10.mx/Heredity/Hurst_partial_1900.html
“When several hybrids from the same pair of species are compared together, this variation of the parts, of “Partial Prepotency,” as I propose to call it, becomes even more apparent and more diverse. For example, in three hybrids raised from the same parents, in the first, the pollen-parent may predominate in form in a certain part; in the second, the seed-parent may prevail in that part; while in the third, that part may be fairly intermediate between both parents; while in regard to colour, these conditions may be exactly reversed. But this only includes one part of the hybrid, and the same law applies equally to every one of the parts so that when the changes are rung on twenty or more different parts by the two parents in both form and colour, we can well understand the many possibilities of variation in hybrids of the same parentage; and I venture to suggest that this law of Partial Prepotency, founded on actual facts observed in hybrids of Paphiopedilum, may perhaps throw some light on the question of variation in offspring of the same parents. Yet, notwithstanding this variation in the parts, it is a remarkable fact that in primary hybrids the whole plant taken together is fairly intermediate between the two parents, the balance of power being well maintained in the whole.”

Larry and Karl, I would like to make a comment about your last couple of posts on this subject if I may. Unless I am reading the wording wrong from Larry’s post, he mentions that hips taken late gave a good germination. Is it that the hips in thier natural environs had warm/partial warm stratification and then cold/some cold stratification ?. In the hybridizers attempt to breed roses he/she does “this” when it suits them, not when it would ‘normally’ suit the plant/nature. Crossing and hip collection might have to be done relative to the type/style of roses that are being used. These are only my immature thoughts. The only thing that gives me more thoughts along these lines is I have ‘Bonica’ growing here and we get hips which have germinated underneath around now, our mid/late Autumn your fall. This might be just a coincidence.

I don’t find significant differences in germination rates between fall-harvested/stratified seeds and spring-harvested seeds that spent the winter in the elements - except for convenience, as David mentioned. The nature-ripened seeds are a bit slower to germinate but do so at the same overall rate. These seeds are all sown outdoors, by the way.

It is necessary to control for moisture to avoid conflating the effects of simple physical disruption due to ice crystals with physiological dormancy. I observed that among seeds from hips that had overwintered on the vine the only viable embryos came from hips or parts of hips that were dry at the time of collection and had apparently dried out before the winter chill.

True, Don - Hips do dry outdoors at some point, which I assume both mitigates moisture damage once winter sets in and eliminates germination inhibition in the pulp, which would presumably be controlled primarily by temperature from that point on. It’s comforting to me that doing nothing can be as effective as obsessive attention to detail.