WI” III I 1|!“le 1 WIN THE. RELATION OF STOCK AND CION EFFECTS TO WINTER HENRY ”V THE APPLE ~. Thesis for the. Degree of M. S. MICHIGAN STATE COLLEGE NOE} E: Doré IQQQ ooh-Id I I.. .u v’dah A .. II. o. 4‘ O .33.: i... _....v”..n.. .. .. .;....§..:u.?£..m. 3%... parts“? . . .. , .. .,. x. ..rn.‘........ . ‘ j; ‘ .T? (uvwrvi.§ow«o.!:‘. .u. ,2; ‘. ‘ . ' .c .. Lou! , ‘ . . 1. .. l . .IULBO'VVOILQM .l... . u. . § {.5‘,‘ | . : -l A, .lo/i 3.40: a 1 .. . . .. . , THE RELATION OF STOCK AND CION EFFECTS T0 WINTER INJURY IN THE APPLE by Noel Etienne Dore “—N A THESIS Submitted to the Graduate School of Michigan State College of Agriculture and Applied Science in partial ful- filment of the requirements for the degree of MASTER OF SCIENCE Department of Horticulture Way]; 443% //." EMA... 1939 TH E515 TABLE OF CONTENTS I. Introduction II. Origin of Grafting III. Relation of maturity to Hardiness IV. Distinction Between Hardiness of the Whole Tree and Hardiness of the Cion Itself V. Review of Literature, to 1900 a. Stock's communication of its own hardiness b. Gain of vigor by grafting as affecting hardiness c. Earlier maturity as affecting hardiness d. Grafting as affecting hardiness of the tree e. Top working vs. root grafting as affecting hardiness and longevity of the tree f. Effect of cion on hardiness of stock g. Conclusions VI. Feview of Literature, from 1900 a. Double working in relation to winter injury b. Hardiness and reciprocal influence of stock and cion VII. Conclusion VIII. Bibliography .4, 53".“: 15‘" ‘3 42‘5“,ng _- \I I.“ Page 01 10 12 21 27 28 THE RELATION OF STOCK AND CION EFFECTS TO WINTER INJURY IN THE APPLE Introduction A review of reports on winter injuries in any particular section of . the United States and Canada shows that the fruit growers experience, every six to ten years, or more often, what is called a test winter and suffer very serious damage in their orchards. Wide and general damage of the severe test winter of 1955—54 in Eastern Canada and in the Norih~ eastern States will long be remembered by producers who went through that winter. Table I is taken fron the data in "The Apple Crop Production and Distribution for 1955" presented by the Dominion Department of Agriculture. The figures on the commercial production of apples give an idea of the acuteness of the hardiness problem for the apple grower of the Provinces of Quebec and Ontario. Table I. - Commercial Production of Apples in 1000 Barrels 5-year ave. Province 1950—51 1951-52 1952-55 1955—54 1929—55 1954-55 Quebec 129 201 258 507 216 121 Ontario 657 976 918 1,069 896 521 These figures show drastic reductions of 70% and 60.5% below 1955 and 44% and 65.6% below the five year averages for Ontario and Quebec, respectively. When one considers the increasing trend in production in the two Provinces, especially Quebec, the figures are still more significant. -2... After a survey in Eastern Canada, Blair (4) listed in order of their importance the different forms of winter killing found in the orchards, as follows: "1. Trunk and body injury, 2. crotch injury, 5. sunscald, 4. bark splitting, 5. killing of dormant buds, 6. killing bark, 7. black heart, 8. discoloration of the sap wood, and 9. trunk splitting." As stated by Blair, the most prevalent injury was to the trunk and at the crotches. More specifically, "the part of the trunk between the snow line and a few feet up the branches was responsible for the majority of the tree losses." This suggests that a substitution of hardy tissues by means of a hardy stock at those weak spots would have helped to overcome the permanent damage to a great extent. This! idea of double working and top grafting to provide hardy trunk or scaffold is by no means new, as it was a common practice in Iowa as earLy as 1870. Budd (ll) of Iowa in 1885 wrote: "While top working has been discussed from a theoretical standpoint, and some half doubtful evidence for many years, we can now, I believe, lay it down as an accepted doctrine that top working varieties as hardy as Willow on stocks not subject to stem injury will add to the longevity and productiveness of our orchard trees. The selection of best stock should now have our special attention." Six years earlier (1879) Budd (9) is reported as saying that the question of influence of stock on cion has been extensiver discussed by practical fruit growers and even respectable biologists for the past hundred years. He considers that the main utility of the hardy stocks for top working is to enable the less hardy cion to endure the test winters of Iowa. Bradford and Cardinell (7) report that, in Michigan, even in the early fifties, few growers and nurserymen realized the possibility - 5 - offered by working the tender varieties into frameworks formed of varieties known to be hardy. The possibility of growing semi—hardy varieties of apple farther North than their normal range by means of suitable hardy rootstocks and intermediate stock, may be considered as the practical application associated with "the relation of stock and cion effects to winter injury". A review of literature is undertaken on the relation of stock and cion to cold hardiness. Attention will be given to the evidence or simple observations tending to show the reality or non-existence of the influence of the rootstock or intermediate stock, or both, on the hardi— ness of the whole tree, or of the cion alone. The reciprocal effect of the cion on the resistance to cold of the stock or stocks will receive due attention. The purpose of this review, however, is not so much to offer an explanation of the phenomenon, as to assemble and present what information is already available, in the hope that it may prove helpful in future investigations of the problem of stock and cion as related to hardiness in fruit trees, especially apples. Origin of Grafting The actual time of the beginning of grafting is unknown. As an art, it appears to be of the remotest antiquity. From history, as Daniel (20) reports, one learns that this art was known by the Chinese 6000 years B. C. The Phoenecians, the Carthagenians, the Arabs, the Greeks and the Romans employed our present methods of grafting, even many of its aspects which are considered as recent discoveries. One may suppose that with the practice of the art of grafting the human mind, always inquisitive, must have wondered about the influence - 4 - of the stock on the cion. It is also conceivable that those people who were near the thermal limits of certain species or varieties of fruits questioned themselves as to whether it would be possible to grow on their own land, in their own regions, those desirable fruits which were flourishing in the gardens of their foreign neighbors who were enjoying a more temperate climate. That they took recourse to the art of graft— ing, hoping to produce the fruits they desired, is likely. The Ancients, as the Moderns, studied the nature of the association between the stock and the cion, but the discussions were metaphysical rather than scientific until and during the Middle Ages. It is just recently that the question of the reciprocal influence of stock and cion began to be systematically studied by the botanists and horticulturists. One must not be surprised then to see in a review of literature the divergent opinions held on the subject and "a fortiori", when one remembers that many of the observations reported were not made by men with scientific training and the ability to make the results of their work very reliable. In many cases at least, on account of the ignorance or carelessness of the observers, the effects attributed to grafting or stock must be explained by other causes. This is more likely to happen when one deals with the hardiness of the fruit trees which can be affected by so many factors outside the possible effect of the stock. Relation of Maturity to Hardiness By far the most important factor determining whether a tree is injured at a given temperature is the degree of maturity of its tissues. Emerson (27) says "Resistance to cold in trees is due often almost wholly to the habit of early maturity rather than to constitutional hardiness........ Northern trees are constitutionally no hardier than Southern but their - 5 - superior resistance to winter cold was due to their habit of ripening their new growth perfectly in the fall." As hardiness,within certain limits, is a function of maturity of tissues it might be inferred that anything which affects maturity of tissue will affect hardiness. Then if a stock can induce earlier maturity it may affect hardiness. Hardi— ness and maturity in fruit trees being so associated, any evidence of the effect of stock on maturity of tissue of cion should receive Special attention. ‘ Distinction Between Hardiness of the Whole Tree and Hardiness of the Cion Itself Very frequently the hardiness of the whole tree, resulting from the symbiosis of stock and cion, is not distinguished from the hardiness of the cion or the grafted variety alone. This lack of precise definition of the problem is responsible for a part of the conflict of evidence or observation that one finds on the effect of stock on hardiness. To make such a distinction is to see that certain contradictory opinions are not real and thus to bring more consistency into the evidence. A grafted tree may be hardier just because a weak part, either the root or the trunk or the crotches, has been replaced by a hardier one, the hardiness of the cion being unchanged. Review of Literature, to 1900 A great majority of the articles written on grafting, before the last century, deal with the methods of doing it and are descriptive of the same. A lesser number of these articles deals with the effect of graft- ing on the vigor and the habit of growth of the tree, on the color, flavor, and maturity of the fruit, on the adaptation of the tree to soil and climate. The effect of the cion on the stock is seldom discussed. _ 5 _ The first mention in English of the influence or use of stock to adapt fruit trees to a colder climate is found in Philip Miller's "Gardener's Dictionary" (55), second edition, 1755. In his article on grafting, he says "....and it is by this Method that many Kinds of Exotick Trees are not only propagated, but also render'd hardy enough to endure the Cold of our Climate in the open Air; for being grafted upon Stocks of the same Sort which are hardy, the Grafts are render'd more capable to endure the Cold; as hath been experimen'd in most of our valuable Fruits now in England, which were formerly transplanted hither from more Southerly Climates, and were at first too impation of our Cold to succeed well abroad, but have been by Budding or Grafting upon more hardy Trees, render'd capable of resisting our severest Cold." Obviously, Miller believed that the hardy stock communicated a part of its own hardiness to the cion: "The Grafts are render'd more capable to endure the Cold." Chronologically the next opinion found on the subject, almost a hundred.years later, is by Knight (44) in his Horticultural Papers: "Many gardeners entertain an opinion that the stock communicates a portion of its own power to bear cold without injury to the species or variety of fruit which is grafted upon it, but I have ample reason to believe that the opinion is wholly erroneous....." Knight, who probably made more experiments and observations in grafting than any man of his time, holds that the hardiness of the stock is not communicable to the cion. This great horticulturist seems to be in contradiction with the idea of Miller and, as he himself says, with many gardeners of his time. Before a discussion of these two divergent opinions, it is worth while to note the various opinions which have been held, and the evidence _ 7 _ on which they have been based, prior to the beginning of this century. For the sake of clarity, these opinions may be placed in two main groups: Those holding that the cion tissues might be influenced in their hardiness by the stock, and those who hold that the stock can not have such an influence. Among the former some claim an effect of stock on the hardiness of the cion, due to communication of its own hardiness or to some change in the habit of growth as increase in vigor, dwarfing effect, or to earlier maturity of the cion wood. ~Stock's communication of its own hardiness That the stock could communicate a part of its own hardiness to the cion, no clear direct opinion of this sort was found. On the other hand, Fuller, Hovey, Budd, Lathrop and Lindley, as we shall see later, declare, like Knight, cited above, that there is not such an influence. Gain in vigor by grafting as affectingghardiness As the result of some observations upon young trees in the Arnold Arboretum of Cambridge, Massachusetts, Jack (45) associates a gain in hardiness in grafted Oak and grafted Ash to an increase' in vigor brought about by the stock. "The dwarf Qpercus geoggiana upon its own root grew more slowly and was less hardy than when grafted on _Qgercus robur. The stocks in this case increased in diameter more rapidly than the grafts. Fraxinus anomala, a southwestern species, also grew more rapidly and was more hardy when grafted upon Fraxinus americana than when grown as seedlings." Wetherell (66) claims that Magnolia glauca, when grafted on Magnolia acuminata, will reach three times its ordinary size and be more hardy. However, Hovey (42) admits the increase in size gained by various other magnolias when grafted on Magnolia acuminata but denies _ 8 _ that there is any gain in hardiness thereby. Earlier maturity as affecting hardiness As we shall see later, it has not been proved yet that the stock may'increase the hardiness of the cion tissue except in so far as it might affect the habit of growth or more especially by hastening the maturity of the wood of the cion. Fuller (52) said "the hardiness of a tree is but slightly changed by the stock except as its growth is influenced to mature early or late in the season." Gideon (55) claims that the crab apple and hardy hybrids make a brief vigorous growth and as they naturally ripen their wood early in the fall, the tendency is for them to force the top to ripen early enough to avoid injury. With the peach, Matthews (55) states that it does better on plum stocks than on peach stocks because it seems to ripen earlier and stand the winter better. An unusual experience worth quoting is reported in 1885 by Cotta (19): "During the last four years I have top worked perhaps five or six thousand trees in nursery, using only ironclads for stems. The results have been surprising.....They all made a fine growth the follow- ing summer and passed the winter of 1882—1885 in first class condition, not a bud being killed. Most of them were taken up and sold. They all grew finely the following year, and are all right todanyJanuary, lBBE/fl "Now compare my record of root grafted trees, standing on the same plat, under the same condition of soil, cultivation, etc. in the spring of 1885. "Uninjured - Duchess, Tetofsky, Whitney and the crabs - 9 _ "Considerably injured - Walbridge, Nelson' Sweet, Wealthy, Haas, Shiawassee, Beauty and Fameuse "Severely damaged - Early Harvest, Red Astrachan, Sweet June, Bailey Sweet, ...., Pewaukee and Tolman Sweet "Partly killed — Sweet Pear, Prices Sweet, ...., Ben Davis "All killed — Hill's Red, Colburn, Koshkamong and Willow Twig." Should we attribute all this to the effect of a hardy stock or, partially at least, to a result brought about by the operation of top working and root grafting. Due to the difference in cutting back, the root grafted tree might have responded by a more succulent growth of the cions, which grew later in the season than in the top grafted trees and did not have as much time to mature their wood to the same extent as the top worked trees. They were consequently less hardy and more likely to be injured by the cold weather of the following autumn or early part of the winter. Here again, as in so many cases, a distinction must be made between the effects of the process itself and the effects of the material used in the process. Grafting as affecting the hardiness of the tree It is interesting to cite the opinion of Downing (26). In the 1869 edition of "Fruits and Fruit Trees of America", he says: "A variety of fruit which is found rather tender for a certain climate or a peculiar neighborhood is frequently acclimatized by grafting it on a native stock of very hardy habits." Without going any farther, one might say that Downing was talking about the influence of the stock on the hardiness of the cion, but such does not seem to be the _ 10 - case because on the next page he says: "If our soil or climate is unfavourable we use a stock which is adapted to the soil, or which will by its hardier root endure cold." Thomas (64), as Downing, claims that trees are made hardier by being grafted on hardier stock. Loudon (47) had the idea that a tree with a tender root is made hardier when grafted on a hardy stock. Lindley (46), in "The Theory of Horticulture", wrote: "It is sometime desirable to increase the hardiness of a variety, and grafting or budding appear to produce this effect to a certain extent, not, indeed by the stock communicating to the cion any of its own power of resisting to cold, but by the stock better suited to the soil of latitudes colder than that from which the cion comes and consequently requiring a lower bottom-heat to arouse its excitability." Top workinggvs. root ggafting as affecting_hardiness and lgggevity of the tree In different periodicals and horticultural society reports one finds opinions like the following on the advantages of top working or double working. Hanford (57) of Wisconsin considers: "Baldwin root grafted, tender in nursery, bark splits and top kills back, worked high seems hardy. Esopus Spitzenburg same as Baldwin, Roxbury Russet tender if root worked, stock worked is more hardy." Beal (5) in Michigan believed that Baldwin at least in some cases is more hardy in cold climates when top worked upon a hardy tree. _ 11 - Fluke (SO) in Iowa during a discussion at a meeting of the State Horticultural Society'stated: "I have Stark, top worked in perfect condition, while the root grafted trees are dead." Later (51) he argues that in his experience some tender varieties are rendered more hardy by being grafted upon hardy stocks. Corbett (18) remarked that even in West Virginia top working can be made profitable to increase the longevity of trees. In The Country Gentleman (l) the editor in an article on the effect of grafting says: "It has been found in Iowa, Quebec, Vermont and other semi—artic countries that comparatively tender varieties of apple (.......) can be grown from 100 to 200 miles further North when top grafted on such 'ironclads' as Hibernal, Duchess or even Tolman." Budd (10) of Iowa found that Jonathan and Dominie will do well on hardy early maturing stocks like Gros Pommier and Duchess but fail when root grafted. Powell (58) of Delaware in an early bulletin of the Station claims that top working provides a healthy strong trunk for all varieties, corrects the poor growth of some and overcomes the tenderness of others in the far North. While holding that top working is advantageous, the following authors consider that the top is not made hardier by a hardy stock. Budd (12) of Iowa says that there is no proof that a tree which is liable to injury in Iowa can be made hardier by grafting it on a hardier intermediate stock, but that top working is of advantage in more than one way. Lathrop (45) in Iowa, after 50 years of experience in top working in his orchard, declares that top working will not make truLy tender varieties hardy, and gives as an example a variety killed back to the stock which remained perfect. To the opinion of Knight (44), Lindley (46), Budd (12), Lathrop (45), holding that the stock does not affect the hardi- ness of the top,we might add the opinion of Hovey (41) of Massachusetts, who said: "If trees could be made hardier by grafting on hardy stocks, that would be a very important point; but the idea of acclimation by this means is utopian." Effect of cion on hardiness of stock The influence of the cion on the stock has always received less attention than the reverse effect of stock on cion. One may suppose that the lack of interest in the stock for itself explains, at least in part, this fact. The stock gets its importance in fruit trees and vines only in so far as it helps the top to produce good fruits in quantity and year after year. Nevertheless few instances of the action of cion on hardiness of stock have been recorded. It is interesting to note that Budd (12), who held that the top is not made hardier upon a hardy stock, found that a tender cion affects the hardiness of the stock and states (15): "With the one thousand or more Gros Pommier stocks used, the great lesson is that the relative hardiness of the variety used in top working affected the hardiness of the stock." With roses, Carriere (16) reports an interesting experience _ 13 _ in France. In one field out of many thousand roses of different sorts, only three varieties were not winter killed. Not a single individual of these three varieties was injured either in the stock or the top. Of the other varieties, except for a few rare individuals, they were completely killed and their stock too. Ward (65),following the severe winter of 1890-91 in France, showed that hardy stocks top worked with cions of tender varieties, such as the tea and Bourbon roses, were killed; and in no instance were such top worked stocks as hardy as the ungrafted hardy types. Conclusions In the light of the information available at the end of the last century, the following conclusions seem to be justified, in an attempt to reconcile the different opinions. A hardy stock may increase the hardiness of the cion, at least in certain cases. The hardy stock, however, does not appear to transfer a part of its own hardiness as a specific quality of its own, but affects the hardiness of the cion indirectly by affecting its habit of growth, that is to say, by increasing the vigor of the top in certain cases, or inducing an earlier maturity in varieties which on their own roots are liable to grow too late in the season. On the other hand, while scarcely noticed, the stock may be increased or decreased in hardiness, at least in certain cases, by a more or else less hardy top. Before ending this review of literature of the last century, it is interesting to note what Bailey (2) wrote of the reciprocal influence of stock and cion in the 1900 edition of his "Cyclopedia - 14 - of Horticulture": "A fourth general office of grafting is to adapt plants to adverse soil or climates..... The practice in Russia of working the apple on roots of Siberian Crab is an example of an effort to make a plant better able to withstand a very severe climate. "In common practice, the effect of the stock on the cion is rather more a mechanical or physical one than physiological or chemical. The influences are very largely those which are associated with greater or less growth, as a rule each part of a combined plant the stock and the cion maintain its individuality. There are certain cases, however, in which the cion seems to partake of the nature of the stock; and others in which the stock partakes of the nature of the .cion." Review of Literature From 1900 The end of the first decade of the 20th Century marked the beginning of a new trend in horticultural research in the United States. Its characteristics have become defined as the employment of laboratory techniques, the use of statistical methods for experimental planning and the analysis of data, and thirdly, a cooperative outlook with reSpect to some of the other sciences, particularly Botany, Chemistry, Soils and Climatology. The result, thus far, has been a measure of progress towards the more fundamental solution of certain major problems in Horticulture that was impossible in the preceding century, with its narrower range of scientific perspective and equipment. With the help of new discoveries, better methods, new tech- niques and greater cooperation, and in the light of the evidence -15- accumulated today, the conclusions drawn at the end of the 19th Century could be changed or at least modified. However, before considering the present ideas on the reciprocal influence of stock and cion on hardiness, it is interesting to note what has been in the last four decades the opinion on the practicability of top working or double working as a means of growing less hardy varieties farther North than their normal limit. Double working in relation to winter injury We already have seen, in the first Section, that topworking or double working on hardy stock is not a new practice in the orchard or the nursery. As it is understood that eXperiences in one part of the country can be applied to another part only after careful trials have been carried out, the idea on the practicability and value of growing otherwise tender varieties on hardy stock will be considered here with respect to the different sections of the country. In Eastern Canada, where hardiness in fruit trees is of the highest importance, Macoun (49) reports: "We have done more or less top grafting here for the past 17 years. I have been interested in it merely from the standpoint of increasing the hardiness of trees which grown as standard trees were not satis- factory. After trying a great many varieties, about 90, I found that the stock did not make the graft sufficiently hardy to with- stand severe winters, so I have come to the conclusion that as far as increasing hardiness is concerned, while top grafting may make some difference, there is not enough to warrant this method." -16.. About 25 years later, conversely to fiacoun (49), Blair (4) does not hesitate to declare that the method of building a hardy framework by double working has been too long overlooked by fruit growers of Canada. He goes as far as to say that this practice is of as great or greater importance than the question of varieties. Blair made the above assertions after an extensive survey on the damage caused by winter injury to fruit trees in Eastern Canada. He also believes that it would pay, in the long run, to plant only trees with hardy framework. The procedure for building such a tree is given in detail: Only seed of hardy Russian varieties are used; the varieties Antonovka, Anis or Charlamoff are used as intermediate stock and the final process is budding at about 18 inches on the main scaffold branches with fairLy hardy standard varieties. From Eastern Canada, crossing to the Northeastern States, we find Potter (57) in New Hampshire who admits that much of the trunk and crotch injuries can be avoided by t0p working certain tender varieties on trees of very hardy sorts. Potter also con- siders that the procedure "involves considerable expense and greatly delays profitable production in the orchard." Unlike Blair (4), however, he does not go as far as to recommend that practice. In a recent bulletin MacDaniels(48) gives preventive and remedial methods for injuries to fruit trees in the orchards of New York, but he does not include top working or double working on hardy framework. Blake (6) in New Jersey says that it would be desirable to _ 17 - have varieties of apple like Cravenstein, Tompkins King, Grimes and Baldwin propagated on a late starting stock such as Northern Spy forming at least a few inches of the trunk. After the severe winter of 1955-56 an extensive survey on winter injuries was made in Ohio. Havis and Lewis (38), like Selby (60) 20 years earlier, found promising, for Ohio, the method of top working on hardy framework. As, at that time, only few orchards were top worked on such hardy stock as Hibernal, Haas and Virginia Crab, they are not as yet able to give definite recommendations. Bradford and Cardinell (7) in Michigan are very explicit: "There can be but little doubt that many orchards which have long ceased to be profitable or have disappeared would still be pro- ductive had they been composed of double worked trees." Rhode Island Greening, Grimes Golden, King, Hubbardston, Cravenstein and Stayman Winesap should be double worked, they say. As to the stocks to work on the recommendations are given with reserva- tions: Duchess and Hibernal "should support such varieties as Vagener and Grimes Golden" but they hesitate to recommend them for "free growers like Greening" stating that probably Fameuse would do better. Tolman Sweet and Astrachan, on account of their susceptibility to fire blight, are disqualified as hardy stocks. In Indiana flcClintock (54) found Virginia Crab most promising as a hardy stock, especially for Grimes Golden and Ben Davis. Delicious proved to be too tender to be used as an intermediate stock. _ 18 _ Much work has been done in Iowa to investigate the possible advantages of double working or top working in the apple orchard. Holland (40) suggests a more extensive use of top working "to make certain varieties better able to withstand unfavorable climate." He reports that Jonathan, Grimes Golden and Delicious top grafted on Hibernal and Virginia Crab are very successful "in Delaware and Clayton Counties" considerably North of their usual limit. Maney and Plagge (50) write: "In Iowa and most other states of the Upper Mississippi Valley as well as Canada, where winter injury is the limiting factor of successful orcharding, topworking on hardy stocks is practical as a method for increasing the hardiness of the best commercial varieties like Jonathan, Grimes and Delicious." To show the beneficial effect of a hardy intermediate stock, they give the following table: Table II. - Record of Topworked Apple Trees in W. R. Campbell Orchard Woodbine, Iowa. Orchard Planted 1895-94. Trees alive Trees alive rPercentage of Variety and Stock 1914 1954 trees alive 1954 Grimes on French Crab 72 5 4.0 Grimes on Virginia Crab 22 16 72.0 Grimes on Haas 155 52 58.0 Grimes on Sheriff 9 7 77.0 Gano on French Crab 84 5 5.5 Gano on Sheriff 57 26 70.0 Sheriff on French Crab 108 66 61.0 Jonathan on French Crab 115 50 26.0 Jonathan on Virginia Crab 15 7 54.0 Jonathan on Haas 44 56 82.0 Jonathan on Sheriff 2 2 100.0 _ 19 - These records show clearly that the length of life of the trees of the several varieties used has been increased by the three hardy stocks used, namely, Virginia Crab, Haas and Sheriff. Maney (51)again reports of 6-year-old trees of Jonathan and Turley on French Crab alternating with double worked on Hibernal. After the severe winter of 1955—56 he found the topworked trees "outstanding in their freedom from winter injury." Understanding, "the fact that a certain stock may be ideal for one variety is no indication that it might do well with another", Maney (52) made a survey to secure some information as to the compatibility of variety and stock combinations for double working. For 45 varieties of apples he gives a "list of stocks used successfully in top working." The increasing interest in top working for hardiness lead the Iowa Station to issue a bulletin entitled "Top Working on Hardy Stocks to Produce Long-Lived Apple Orchards". A procedure is given to form trees with hardy framework. In Missouri Talbert (65) informs us that some top working is done on Minkler, Arkansas, Hibernal, Virginia Crab and others to replace the weak crotch structure and avoid the crown troubles of Grimes and the standard varieties, but not so much for the hardiness of the same parts. In Wisconsin, Minnesota and the Dakotas, many of the semi-hardy but desirable varieties are recommended only as top worked trees on hardy intermediate stocks as Hibernal and Virginia Crab. In the Pacific Northwest, the search for the best stocks for double working is also going on. In Oregon at the Hood River Station the following varieties have been found promising as intermediate stocks: - 20 - Astrachan, Arkansas, Cortland, Scott Winter, Hibernal, Antonovka, Oskakoff and also McIntosh (17). It is worth noting that Virginia Crab has not proven hardy enough (8). The use of double working in an attempt to overcome some winter injuries has also received some attention in central Europe. In Poland, Filewicz (29) stresses the value of double working. Antonovka is the stock extensively used an an intermediate. The recommendations of Filewicz differ from all that we have noted so far in one point — one or two braches of the hardy intermediate stock are allowed to grow and he claims that these increase the resistance of the tender variety: "The trees .... on hardy trunks do not suffer from frost, and because of the symbiosis of tender variety with a hardy one they neither need bridge grafting nor invigorating." It would seem interesting, before ending this chapter, to have a summary of the work that is going on in Canada to find the best adapted rootstocks and intermediate stocks to meet the requirements of different regions. At Ottawa, Ontario, they are comparing the single worked trees with double worked trees on hardy Russian varieties, namely, Hibernal, Antonovka, Anis and Charlamoff. These are all propa— gated on uniform hardy rootstock, "which is considered to be quite homozygous." Finally these are worked over to McIntosh and Fameuse by inserting buds 18 to 24 inches from the trunk on the main scaffold limb. This experiment divided into six blocks is planned to permit the use of the best methods of statistical analysis (5). At Cap Rouge, Quebec, a similar experiment is going on. The commercial varieties, McIntosh, Lawfam and Lobo, are top grafted on Antonovka, Hibernal, Charlamoff and Patten Greening as intermediate _ 21 _ stocks. The rootstock is Anis. The experiment is divided into six blocks also (59). In Western Canada, some work is also being carried on along the same line at Sommerland, B. C. From this inquiry into the practice of double working on hardy intermediate stocks, one sees that new interest has recently arisen about the practical advantages in the method to grow hardier, healthier and longer lived trees. This beneficial effect is expected to be accomplished not so much by the influence of the stock on the hardiness of the top, as to fit a top with a hardier trunk or hardier crotches and render the tree more able to struggle successfully against cold injuries. In the next chapter we shall see if the top, in the grafted tree, partakes of the hardiness of the stock or may be influenced in its hardiness some other way by grafting. Hardiness and reciprocal influence of stock and cion It is a general belief that the use of a hardy stock is an effective means of making a tree better able to withstand cold weather. The hardy stock may be beneficial merely by substitution of tender parts with hardier ones, by the building up of a hardier root, trunk or crotches. But the question arises - does the hardier stock exercise a beneficial influence on the hardiness of the cion tissue of the variety grafted upon it? We already have noted in the first Section a few observations on this point; we shall note a few more that have more recently been reported. At Ottawa, Canada, 90 varieties of apples were top grafted on hardy stocks to see if they could be made hardy, but practically all were killed back to the stock. The cions were six years old or less. -22.. Macoun (49) concluded that if the top varieties are made hardier it is not enough to enable them to withstand very severe winters. It may be said here that examples by thousands could be given where tender grafted varieties have been killed back to the stock. These negative results, though, should not overshadow the few controlled cases where the cion itself may have been rendered more hardy by grafting on a hardy stock which did influence its grafted associate. It is interesting to quote the opinion of Culley (56) who declares: "Without doubt both hardiness and stronger growth are trans— mitted to an extent, from stocks having those characteristics. But I have yet to note that these are in any sense permanently transferred to the top or that the fruit is affected other than environment would account for." With Macoun (49) tender tops were not made, by the stock, hardy enough to endure the test winter of 1905-04. It is supposable, though, that some tops may have been slightly hardier but not enough to make them endure such a test winter; so it may be said with Hedrick (59): "It would be too much to say that hardiness as an inherent quality is transmitted from stock to cion, but it is very certain that increase in vigor imparted by some stocks gives better hardiness to grafted plants. In the case of late growing apples worked on Hibernal or Oldenberg stock greater hardiness results because the cion matures earlier than it would upon its own roots and is therefore better able to withstand the winter. Again,.slight obstructions to the sap flow imposed more or less by union of different tissues of stock and cion may cause the cion to ripen earlier and this bring about greater hardiness." _ 25 _ Dorsey (24) in 1918, using the degree of browning of wood as an index of winter injury, undertook one of the first systematic laboratory studies of the reciprocal influence of stock and cion on hardiness. The degree of injury was determined by cutting, into selected limbs of one to one—and~a—half inches in diameter, three to five annular rings deep with a sharp knife or hatchet. Dorsey considered that the browning, varying with the severity of the winter and with the hardiness of a variety, was "a most sensible index of the winter injury." The conclusions he arrived at were as follows: "The hardiness of the cion is independent of that of the stock and if a hardy stock does have an influence upon the hardiness of the cion, it is not sufficient to prevent it from being injured by low temperatures." Dorsey states elsewhere (25): "Fundamentally the stock does not increase the hardiness of the cion except in so far as maturity and nutrition may have a bearing upon hardiness." Working on the same line Carrick (15) undertook an experiment to find out whether the cions of varieties of different hardiness did not influence the hardiness of the stock roots grafted under them. Six hundred forty piece-root grafts of four different varieties were made on stocks of long rooted American seedlings cut into four parts. Each variety was grafted on each of the four cuts. As an index of injury in the roots, the percentage of cells killed in the cambium phloem and Cortex region was taken. No "constant nor considerable differences in hardiness" were noted in the hardiness of the one—year roots of the stock grafted under the four different varieties. It was also shown that there is no significant variation in the hardiness of the grafted and ungrafted seedling stocks. - 24 - Gardner, Bradford and Hooker, in their book, "The Fundamentals of Fruit Production", write: ".....that hardiness of the stock increases the hardiness of the cion is not shown conclusively by any evidence available. It is conceivable that an early maturing stock might influence the top slightly in the same direction but any influence of this character is comparatively insignificant." (55). In another chapter, it is said also: "It is conceivable that a stock may in itself be hardy but through the congeniality of the graft it may actually diminish the hardiness of the cion." (54). Potter (56),in accordance with results reported by Carrick (15), cited above, concluded from his own experiments that cions of varieties of different hardiness do not influence the hardiness of the stock roots which are grafted under them. Garrick had worked with one-year- old grafts while Potter reached the same conclusions with threefiyear— old grafts. Potter states that apparently no compounds which influence hardiness are translocated from the stock to the cion. It should be pointed out here that neither Potter nor Garrick worked with clonal rootstocks. The variability due to the different genetical make-up of seedling rootstocks may possibly be responsible, at least in part, for their failure to find an influence which, if it exists, must be of small magnitude. The observations of Filewicz (28), as we shall see, would seem to show that the influence of the two or more components of a grafted tree, on one another, is dependent on the age of the union or unions. Filewicz states that tender varieties could be made more resistant by top graft— ing with hardier varieties while hardiness could also be diminished by tender top varieties. His findings are based on extensive observations -25.. in the orchards for many years and more especially after the unusually severe winter of 1928-29 which caused so much damage to fruit trees in Central Europe. He sums up as follows the observations he made on Conlon's Reinette, a variety grown there for its fruits as well as an intermediate stock: "1. when not top grafted a large percentage died of frost; 2. after cutting it offers little resistance against frost for many years. All trees grafted since 1925 with varieties of medium or little resistance died after the winter of 1928-29; 3. grafted with a very resistant variety (Antonovka) in 1924 the trees became resistant in five years, and 4. grafted with a fairly resistant variety (Broiken) only after ten years." It has been shown above that Garrick (15) as well as Potter (56) failed to find any consistent augmentation of hardiness, in the root- stocks, attributable to the cion grafted on. They were working with one-year and threeayear-old grafted material, respectively. These were relatively short unions when we consider that with Filewicz (28) it took five years for a very resistant variety to make its intermediate stock hardy and as long as 10 years for a fairly resistant variety to accomplish the same. One many infer, though, that an intermediate stock can possibly react in a different way, as a rootstock does, and conceivably can be slower or faster to respond to the action of the cion. Filewicz (29) again claims that a tender top is made more hardy if a few grafts of a hardy variety are grown in it, or if one or two branches of the hardy intermediate stock are allowed to bear foliage. Would these observations corroborate the views of Burbidge (14) who wrote in the last century: "Do we not rob the stock of a deal of its power to ameliorate _ 25 _ the scion when we denude it of all its own leaves?" I should think that this question, of the effect of a not completely defoliated stock as compared to a completely defoliated one, should receive more attention than has been devoted to it so far. The hard test winter of 1955-54 gave to Blair (4) a very good opportunity to analyze statistically winter data. Thirteen different varieties of apples grafted on six different rootstocks were growing in six randomized blocks. The unions were four years old. The analysis of variance of the injury measured after a given system of pointing was done. Not even in a single instance was a significant difference between any of the rootstocks as imparting hardiness to any double worked variety found. The examination of the literature so far would seem to point out the need of a practical and sensible test to be applied to the solution of the problem. Recently, the exosmosis method of measuring cold resistance devised by Dexter (21) (22) (25), and tested for apple tissues by Swingle (62), was tried on the problem by Stuart (61). In order to avoid variability in seedlings due to different genetical make—up, clonal rootstocks were used. Contrary to Potter (56) and Garrick (15), Stuart found that the cion can appreciably affect the hardiness of the rootstock, but it is worth noting that the direction of this influence could not be predicted from the hardiness of the cion variety. As to the effect of stock on the hardiness of the cion, the results were different: no significant appreciable difference was found. The material used in this experiment was only one growing—season old after budding. _ 27 _ Conclusion After this review of literature dealing with the subject, one sees that the reciprocal influence of stock and cion on hardiness is not a problem that has ceased to be controversial. Little conclu- sive information is as yet available and the problem stays an open one for investigation and research. Probably very sensitive and accurate method of testing hardiness is necessary before there can be a closer approach to its solution. Parenthetically, the method based on electrical conductivity used by Stuart (61) would seem to be promising. Also in an endeavor to minimize the difference necessary to be significant, any eXperiment on the subject should be previously planned to permit the use of the best and more refined methods of statistical analysis. Finally, variation in the material should be controlled as much as feasible, so clonal rootstocks should be used, even in an experiment to test the influence of the intermediate stock on the cion. (5) (6) (7) (8) (9) (10) (ll) ((12) (15) (14), (15) (16) (17) (18) (19) - 28 - Bibliography .......... - The Effect of Grafting. Country Gentlemen. 64:588. 1899. Bailey, L. H. - The Cyclopedia of Horticulture. New York. 1900. Beal, W. J. - Michigan Board of Agr. Report. p. 204. 1876. Blair, D. S. — Winter Injury to Apple Trees in Eastern Canada. Scientific Agr. 16:8-15. 1955. Blair, D. S. - Correspondence. August, 1958. Blake, M. A. - Winter Injury to Fruit Trees in New Jersey. Am. Soc. Hort. Sci. Proc. 15:24-25. 1917. Bradford, F. C. and Cardinell, H. A. — Eighty Winters in Michigan Orchards. Mich. Agr. Exp. Sta. Spec. Bul. 149. 1926. Brown, G. G. Correspondence. September, 1958. Budd, J. L. - Influence of Stock on Scion. Iowa Hort. Soc. Trans. 14:464. 1879. Budd, J. L. Mass. jort. Soc. 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