Are infrastipular prickles really modified stipules

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Are infrastipular prickles really modified stipules

Postby Karl K » Tue Dec 20, 2016 11:51 am

While looking into shoot tip abortion, I came across a fascinating book, 'On Buds and Stipules' by Sir John Lubbock.
https://archive.org/stream/onbudsstipul ... 1/mode/2up

On page 188 the author discussed Rosa persica:
"The stem is prickly, and the frequent occurrence of the prickles, sometimes in pairs, at the base of the leaf has led to their description as stipules; for instance, by Boissier in his great ‘Flora Orientalis’."

Lubbock did not offer his opinion on the accuracy of this observation. But it did get me to thinking about infrastipular prickles in general. Lubbock gave examples of prickles that are undoubtedly stipules, such as those of Robinia pseudoacacia.
http://www.rnr.lsu.edu/plantid/species/ ... 3a_web.jpg

Now, compare those to the infrastipular stipules of Rosa woodsii.
https://www.minnesotawildflowers.info/u ... i-15-2.jpg

The relationship of stipules to leaflets is sometimes revealed in hybrids. For example, the orange has simple leaves. The trifoliate orange (Poncirus trifoliata) appears to have three leaflets, but technically the side-leaflets are stipules. But a hybrid of trifoliata and the Thompson Navel orange shows many 5 foliate leaves.
http://bulbnrose.x10.mx/Heredity/Swingl ... 911_07.jpg

I am interested in the occurrence of infrastipular prickles in rose hybrids, and whether these are correlated (linked) with other traits of the parent that provided them.
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Re: Are infrastipular prickles really modified stipules

Postby Don » Sat Dec 24, 2016 2:33 pm

It is an interesting question and your suggested approach using hybridity seems like a really good one but wonder if ploidy differentials might interfere. My thought experiment would use moyesii, which has huge and long stipules attached along the leaf stem but, being heavily polyploid, might it not yield both stipules and prickles?

Actually, I wonder if I have already done the experiment. I will need to have a look in the spring.
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Re: Are infrastipular prickles really modified stipules

Postby Karl K » Sat Dec 24, 2016 7:49 pm

Infrastipular stipules seem to be a long-established character of Rosa, whether habitual or occasional. Getting these to revert to a more "traditional" sort of stipule might require intergeneric hybridization - like the hybrid of Poncirus and Citrus seemingly reverting to an ancestral form with pinnate leaves.

And on that possibility, I recently found a very suggestive paper by O. F. Cook, 'Jointed leaves of Amygdalaceae.' (1912).
http://bulbnrose.x10.mx/Heredity/CookLeaves1912.html

Amygdalaceae is the old name for Prunus. Cook pointed out that the joint at the base of the leaf, and nectaries on the leaves of some apricots, are evidence that these species are descended from ancestors with compound leaves.
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Re: Are infrastipular prickles really modified stipules

Postby Karl K » Mon Feb 13, 2017 11:10 am

The Gardeners' Monthly and Horticulturist, 29: 376-377 (Dec 1887)
On The Stipules Of Magnolia Frazeri.
Thomas Meehan
"In many species of roses, especially in Rosa Kamtschatica and Rosa cinnamomea, the stipules could be noted increasing, and the size of the leaf blade diminishing on the branch as it approached infloresence. Often the tips of the sepals would develop to minute leaf blades; and in a few instances he had seen the same appendages on abnormal petals. Often the stipules, especially in Rosa Kamtschatica, would have the red colors of the petals, when at the nodes, immediately below the axis from which the peduncle proceeded. There could be no possible doubt in the minds of those who would carefully compare, and watch for occasional aberrations, that the petals of the rose were rather transformed petals [sic: I think this should be "transformed stipules"] than complete leaves."
http://bulbnrose.x10.mx/Heredity/Meehan ... s1887.html
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Re: Are infrastipular prickles really modified stipules

Postby Don » Mon Feb 13, 2017 6:19 pm

>> the petals of the rose were rather transformed..transformed stipules..than complete leaves.

Hmmm. I would think that this question has already been answered by the folks who study inflorescence. Does anybody here from academia know if petals and stipules are the same bird?
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Re: Are infrastipular prickles really modified stipules

Postby Karl K » Tue Feb 14, 2017 9:27 am

Don wrote:>> the petals of the rose were rather transformed..transformed stipules..than complete leaves.

Hmmm. I would think that this question has already been answered by the folks who study inflorescence. Does anybody here from academia know if petals and stipules are the same bird?

I would be interested to learn of any research subsequent to Meehan's.

BTW, the Bot. Reg. illustration of Rosa ferox shows the red pigment in the stipules immediately below the flower. At this point in my research, R. ferox and R. kamchatica seem to be selections of the same, small-leafed species.
http://bulbnrose.x10.mx/Roses/Rose_Pict ... ferox.html

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Re: Are infrastipular prickles really modified stipules

Postby Karl K » Wed Feb 15, 2017 9:37 am

A further report from Meehan on stipules and petals:

Proceedings of the Academy of Natural Sciences of Philadelphia 41: 53-66 (1889)
Contributions to the Life Histories of Plants. No. IV
Thomas Meehan
Clear as it is to the mind that when carefully traced, the petal of a rose is formed of an enlarged stipule, and not of a fully planned leaf, the positive evidence is not furnished as freely as in the case of the sepal, but specimens of Rosa humilis, sent to me in 1883 by Miss Jennie E. Whiteside, of Harmonsburg, Pennsylvania, give an excellent illustration. This form has been figured and described by Mr. Sereno Watson in the Garden and Forest for February 13th, 1889 as Rosa humilis, var triloba. The trilobed petal is simply a case in which the usual stipule forming the petal of the rose, has again had its normal growth accelerated towards a perfect leaf. The central lobe is in fact no more than a dilated petiole, with the stipule represented by the two lateral lobes, in its normal position at its base. The same process from the total arrestation of petiole and leaf blade to the abnormal dilation of the stipule to form the petal, can be traced in magnolia, as made plain in the paper above cited.
https://books.google.com/books?id=dHkOA ... &q&f=false


Image
Rosa humilis var triloba
http://bulbnrose.x10.mx/Roses/Rose_Pict ... iloba.html
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Re: Are infrastipular prickles really modified stipules

Postby Karl K » Thu Feb 23, 2017 4:12 pm

I've been reading about extrafloral nectaries and how they attract ants that devour sucking insects that bother the same plants. In Guatemala, the Kekchi cotton plays host to keleps (a species of ant) that sip the nectar and dine on boll weavels.

A little more searching turned up this item;

Annual Report of the Board of Regents of the Smithsonian Institution for the Year ending June 30, 1896 pp. 421-422
The biologic relations between plants and ants
Dr. Heim
Associate of the Faculty of Medicine at Paris

Along the edges of the leaves of the Rosa Banksiae are found perifoliary nectaries that attract great numbers of a large black ant (Camponotus pubescens). The presence of these ants preserves the rose from the attacks of a hymenopterous insect (Hylotoma rosae). We owe an interesting experiment upon this subject to Beccari. On a branch of Rosa Banksiae attacked by ants he placed a branch of another rose bush attacked by the larvae of Hylotoma. Incommoded by the ants, these larvae took refuge upon the youngest buds, unprovided as yet with nectaries, and consequently not visited by ants. It is to be remarked that the Banks rosebushes, which are rarely or never attacked by Hylotomas, are destitute of prickles. We may probably admit that there is a correlation between the presence on plants of thorns or prickles and that of leaf-eating insects. Is it not due to the protection given by ants and other sting-bearing hymenoptera that the Banks rosebushes attain the great age that some of them are known to do? We may cite as an instance one of these bushes planted in 1803 by Bopland in the garden of the marine hospital at Toulon, which has a stem a meter in diameter at the base and bears each year from fifty to sixty thousand flowers.

The leaves of peach, apricot, and cherry trees may, as there is reason to suppose, be derived from compound leaves. The nectaries which they carry on the petioles should then have the significance of aborted leaflets filled with sweet stores.

https://books.google.com/books?id=a8DaQ ... &q&f=false

So, here is another reason to breed with Rosa banksiae, while paying attention to which seedlings are most often visited by ants.

And another:
The Biology of Nectaries, p. 193 (1983)

Extrafloral Nectaries: Their Structure and Distribution
Thomas S. Elias
One or more, often paired nectaries of the Flach or Hoch type are present at the distal part of the petiole in many species of Prunus and some species of Rosa. In addition, some members of this genus have branched stipules which are nectariferous.
https://books.google.com/books?id=9ULQh ... ry&f=false


Has anyone seen ladybugs (or ladybird beetles) sipping nectar from the leaves or stipules of roses?
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Re: Are infrastipular prickles really modified stipules

Postby roseseek » Fri Feb 24, 2017 12:49 am

Interesting, thanks, Karl. I have not noticed that.
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Re: Are infrastipular prickles really modified stipules

Postby Karl K » Fri Feb 24, 2017 9:01 am

The statement about a Rosa banksiae with a trunk a meter in diameter seems to be a mistranslation. I found the German text, which states that the trunk was a meter in circumference.

The original source was Beccari's 'Piante ospitatrici ossia piante formicarie della Malesia et della Papuasia,' (1885) So, the association of Rosa banksiae with the black ants apparently was observed in the rose's natural habitat.

However, both the ant and the rose sawfly seem (from what I've been able to find) to be natives of Europe.
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Re: Are infrastipular prickles really modified stipules

Postby Karl K » Sat Feb 25, 2017 9:06 am

Delpino (1899) continued the study of extra-floral nectaries. He stated that Beccari had made his observations in Florence, then added his own from Bologna.

Delpino confirmed Beccari's observations of leaf-nectaries on Rosa banksiae, then added that these are also found on the leaves of R. bracteata, though they are less numerous. Furthermore, in Bracteata the nectaries are active only on the young leaves.

Bullettino dell'Orto botanico della R. Università di Napoli, Volume 1 (1899)
Piante Formicarie (Seguito)
Prof. Federico Delpino
Rosa bracteata. -- L’inspezione attenta del diportarsi delle formiche mi ha rivelato la esistenza dei minuscoli nettarii di questa specie. Invero i suoi vigorosi rampolli scarseggiano di spine, e mancano dei soliti peli glandolosi allontanatori delle formiche. Le foglioline sono serrate, e l’apice dei denti, a vece di essere occupato, come nelle altre specie di rose, da collofori, porta un piccolo nettario mellifluo. Questa secrezione, ch'è poco diuturna, riscontrandosi soltanto nelle foglie giovani, adesca un certo numero di formiche, che dimorano verso le sommità vegetative dei rampolli medesimi, e passano lentamente dalle foglioline d‘ una foglia a quelle di un'altra, visitando metodicamente l‘apice dei denti fogliari. La quantità del miele emanato è di gran lunga inferiore a quella della Rosa Banksiae; ma è anche proporzionalmente minore il numero e la statura delle formiche accorrenti. Prendendo la media di molte osservazioni ho rilevato nella sommità di ogni rampollo la presenza o di una sola formica di una mezzana statura, o di quattro o cinque di piccola.

È notevole cosi in questa che nella Rosa Banksiae la mancanza dei soliti peli glandulari; i quali veramente qui sarebbero un contro senso, perocchè sono allontanatori delle formiche.

https://books.google.com/books?id=mwI0A ... &q&f=false
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Re: Are infrastipular prickles really modified stipules

Postby Karl K » Sun Mar 05, 2017 9:34 am

In reading the early descriptions of Rosa rugosa and its varieties, I began to wonder whether the needle-like prickles (glandular acicles) were derived from trichomes. A quick search turned up this:
Understanding Plant Anatomy, p. 123 (2009)
By S. R. Mishra
"Prickles are classified as trichomes, no matter how massive they may be, whenever it is clear that they arise from the epidermis and are not modifications of any other organ.

As a matter of fact, the large prickles of Rosa and the vascular prickles on the fruit of Horse Chestnut are connected with simple hairs by all gradations of finer prickles."
https://books.google.com/books?id=ZAQAq ... &q&f=false


Trichomes are a fascinating subject on their own. Beal (1878) described and illustrated some of the diversity of structure and function he found among trichomes of various plants.
https://www.jstor.org/stable/2448153?se ... b_contents

And Cannon (1909) compared the trichomes of some hybrids with those of their parents.
http://www.biodiversitylibrary.org/item ... 1/mode/1up

Pondering this information, I remembered a very curious curiosity I read about years ago, Begonia phyllomaniaca. Could there be a connection between the proliferation of small leaves on stems and leaves of this begonia? Yes, according to Smith (1919).
"My observations contradict those of Prillieux and confirm those of Verlot and of Caruel that buds may arise from the ordinary trichomes. They may develop either from the base or the middle of acicular hairs. Such hairs arise from a red tissue, the other parts of the epidermis being green. I have also seen them developing from the base of glandular hairs which are abundant on the young internodes, but they are not restricted to these pairs."
http://bulbnrose.x10.mx/Heredity/Begoni ... a1919.html


Another item I found several years ago is also of some interest. MacDougal (1903) noted that the orchid Cypripedium montanum normally produces two types of trichomes, the pointed and the knobbed.
... the growth of Cypripedium in darkness is characterized by a non-development of the pointed hairs on the leaves, and the excessive development of the glandular trichomes.
http://bulbnrose.x10.mx/Heredity/MacDou ... edium.html
http://www.biodiversitylibrary.org/item ... 2/mode/1up


So, the various types of trichomes (and their derivatives) are largely independent in their heredity and in their environmentally modified expressions.

Which brings me to the Rose matter.

Les Roses, pp. 47-48 (1817)
Redouté
It is from one of our drawings that this Rose [R. kamchatica] has been engraved for the garden of CELS, page and figure 67. By comparing this individual with that which accompanies our description, it will be easily seen that in less than eighteen years, this rose has undergone considerable modifications in the length or density of the spines, and in the shape of the leaflets.
http://bulbnrose.x10.mx/Roses/Rose_Pict ... atica.html


In fact, this Rosa kamchatica had become as "fierce" as R. ferox Lawrance. The above link shows Redouté's two illustrations, along with the one from the Botanical Register.

My suspicion is that the original population of R. kamchatica is polymorphic for a gene (or two) that allow for an extreme development of the glandular acicles, but the trait is suppressed by some environmental condition active in Kamchatka. When various specimens are moved further south, some of them become "fierce" while others don't. This suspicion is not contradicted by Palibin (1899) who noted that Rosa rugosa var. ferox is widely distributed, but is NOT found in Kamtschatka.
https://books.google.com/books?id=53g1A ... &q&f=false

Another example of morphological traits being expressed differently under different environmental conditions was noted by Hurst.
Experiments in Genetics (1925)
Charles Chamberlain Hurst
Chromosomes and characters in Rosa and their significance in the origin of species
In the most complex case studied, in the octoploid species BBCCDDEE (R. acicularis Lindl.), the four double septets seem to work more or less in relays in different parts of the plant at different times and seasons, resulting in a periodic predominance of one septet over another in certain parts of the plant, the general result being more or less a mosaic of the four septets of characters arranged end to end or side by side.
http://bulbnrose.x10.mx/Roses/Hurst/HURST2.HTM
Naturally with four double septets working equally and independently in an octoploid species, only about one-fourth of the characters of each septet can be represented at one time.


One final note on the function of glandular organs:
ACTA AGROBOTANICA, Vol. 67 (4), 2014: 13–24
© The Author(s) 2014 Published by Polish Botanical Society
CHARACTERISTICS OF THE SECRETORY STRUCTURES IN THE FLOWERS OF Rosa rugosa Thunb.
Aneta Sulborska, Elżbieta Weryszko-Chmielewska

A b s t r a c t
Due to the presence of secondary metabolites exhibiting pharmacological activity, the flowers of Rosa rugosa Thunb. have found application in traditional and folk medicine. The essential oil obtained from them is also considered to be a phytoncide. The morphological and anatomical characters of glandular trichomes located on the sepals of R. rugosa were studied by light and scanning electron microscopy. Using histochemical tests, the type of secretion produced in the trichomes was determined and its contents were compared with the secretion produced by the papillae on the petals.
http://agro.icm.edu.pl/agro/element/bwm ... -SM_13.pdf


https://en.wikipedia.org/wiki/Phytoncide

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